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fairly regular arrangement from the region included between the
first and the last pair of true feet. There are nine or ten of them
between each pair of feet. They pass along the ventral wall of
the body, perforating the ventral mass of longitudinal muscles.
On their way they give off nerves which innervate the skin.
Posteriorly the two nerve-cords nearly meet immediately in
front of the generative aperture, and then, bending upwards, fall
into each other dorsally to the rectum. They give off a series
of nerves from their outer borders, which present throughout
the trunk a fairly regular arrangement. From each ganglion
two large nerves (pn) are given off, which, diverging somewhat
from each other, pass into the feet.
From the oesophageal commissures, close to their junction
with the supra-oesophageal ganglia, a nerve arises on each side
which passes to the jaws, and a little in front of this, apparently
from the supra-oesophageal ganglion itself, a second nerve to the
jaws also takes its origin.
The supra-oesophageal ganglia (Fig. 9).are large, somewhat
oval masses, broader in front than behind, completely fused in
the middle, but free at their extremities. Each of them is pro-
longed anteriorly into an antennary nerve, and is continuous
behind with one of the oesophageal commissures. On _ the
ventral surface of each, rather behind the level of the eye, is
placed a hollow protuberance (Fig. 9, d), of which I shall say
more in dealing with the development. About one-third of: the
way back the two large optic nerves take their origin, arising
laterally, but rather from the dorsal surface (Fig. 9). Each of them
joins a large ganglionic mass placed immediately behind the retina.
The histology of the ventral cords and oesophageal commis-
sures is very simple and uniform. They consist of a cord almost
wholly formed of nerve-fibres placed dorsally, and of a ventral
‘layer of ganglion cells.
The Body Wall.
The skin is formed of three layers.
(1) The cuticle.
(2) The epidermis or hypodermis.
(3) The dermis.
The cuticle is a thin layer. The spines, jaws, and claws are
special developments of it. Its surface 1s not, however, smooth,
—
14 PERIPATUS CHAP. —
but is everywhere, with the exception of the perioral region,
raised into minute secondary papillae, which in most instances
bear at their free extremity a somewhat prominent spine. The
whole surface of each of the secondary papillae just described is
in its. turn covered by numerous minute spinous tubercles. |
The epidermis, placed immediately within the cuticle, is
composed of a single layer of cells, which vary, however, a good
deal in size in different regions of the body. The cells excrete
the cuticle, and they stand in a very remarkable relation to the
secondary papillae of the cuticle just described. Each epidermis
cell is in fact placed within one of these secondary papillae, so
that the cuticle of each secondary papilla is the product of a
single epidermis cell. The pigment which gives the characteristic
colour to the skin is deposited in the protoplasm of the outer ends
of the cells in the form of small granules.
At the apex of most, if not all, the primary wart-like papillae
there are present oval aggregations, or masses of epidermis cells,
each such mass being enclosed in a thickish capsule and bearing a
long projecting spine. These structures are probably tactile organs.
In certain regions of the body they are extremely numerous ; more
especially is this the case in the antennae, lips, and oral papillae.
On the ventral surface of the peripheral rings of the thicker
sections of the feet they are also very thickly set and fused together
so as to form a kind of pad (Figs. 6 and 7). In the antennae
they are thickly set side by side on the rings of skin which give
such an Arthropodan appearance to these organs in Peripatus.
The Tracheal System.
The apertures of the tracheal system are placed in the depres-
sions between the papillae or ridges of the skin. Each of them
leads into a tube, which may be called the tracheal pit (Fig. 10),
the walls of which are formed of epithelial cells bounded towards
the lumen of the pit by a very delicate cuticular membrane con-
tinuous with the cuticle covering the surface of the body, The
pits vary somewhat in depth ; the pit figured was about 0°09 min.
It perforates the dermis and terminates in the subjacent muscular
layer.
Internally it expands in the transverse plane and from the
expanded portion the tracheal tubes arise in diverging bundles.
Nuclei similar in character to those in the walls of the tracheal
I TRACHEAL, MUSCULAR AND VASCULAR SYSTEM 15
pit are placed between the tracheae, and similar but slightly more
elongated nuclei are found along the bundles. The tracheae are
minute tubes exhibiting a faint transverse striation which is prob-
ably the indication of a spiral fibre. They appear to branch, but
Fie. 10.—Section through a tracheal
pit and. diverging bundles of
tracheal tubes taken transversely
to the long axis of the body.
(After Balfour.) tr, Tracheae,
showing rudimentary spiral fibre ;
tr.c, cells resembling those lining
the tracheal pits, which occur at
intervals along the course of the
tracheae ; ¢r.o, tracheal stigma ;
tr.p, tracheal pit.
only exceptionally. The tracheal apertures are diffused over the
surface of the body, but are especially developed in certain regions.
The Muscular System.
The general muscular system consists of—(1) the general
wall of the body; (2) the muscles connected with the mouth,
pharynx, and jaws; (3) the muscles of the feet ; (4) the muscles
of the alimentary tract.
The muscular wall of the body is formed of—(1) an external
layer of circular fibres; (2) an internal layer of longitudinal
muscles,
The main muscles of the body are unstriated and divided into
fibres, each invested by a delicate membrane. The muscles of the
jaws alone are transversely striated.
The Vascular System.
The vascular system consists of a dorsal tubular heart with
paired ostia leading into it from the pericardium, of the pericar-
dium, and the various other divisions of the perivisceral cavity
(Fig. 14, D). As in all Arthropoda, the perivisceral cavity is a
haemocoele ; 7.e. it contains blood and forms part of the vascular
system. The heart extends from close to the hind end of the
body to the head.
16 PERIPATUS CHAP.
The Body Cavity.
The body cavity is formed of four compartments—one central,
two lateral, and a pericardial (Fig. 14, D). The former is by far
the largest, and contains the alimentary tract, the generative
organs, and the slime glands. It is lined by a delicate endo-
thelial layer, and is not divided into compartments nor traversed
by muscular fibres. The lateral divisions are much smaller than
the central, and are shut off from it by the inner transverse band —
of muscles. They are almost entirely filled with the nerve-cord
and salivary gland in front and with the nerve-cord alone behind,
and their lumen is broken up by muscular bands. They further
contain the nephridia. They are prolonged into the feet, as is the
embryonic body cavity of most Arthropoda. The pericardium con-
tains a peculiar cellular tissue, probably, as suggested by Moseley,
equivalent to the fat-bodies of insects.
Nephridia.
In Peripatus capensis nephridia are present in all the legs.
In all of them (except the first three) the following parts may
be recognised (Fig. 11) :—
(1) A vesicular portion opening to the exterior on the ventral -
surface of the legs by a narrow. passage.
(2) A coiled portion, which is again subdivided into several
sections.
(3) A section with closely packed nuclei ending by a some-
what enlarged opening. :
(4) The terminal portion, which consists of a thin-walled
vesicle.
The last twelve pairs of these organs are all constructed in a
very similar manner, while the two pairs situated in the fourth
and fifth pairs of legs are considerably larger than those behind,
and are in some respects very differently constituted.
It will be convenient to commence with one of the hinder
nephridia. Such a nephridium from the ninth pair of legs is
represented in Fig. 11. The external opening is placed at the
outer end of a transverse groove at the base of one of the legs,
while the main portion of the organ lies in the body cavity in
the base of the leg, and extends into the trunk to about the level
I : NEPHRIDIA 17
of the outer edge of the nerve-cord of its side. The external
opening (0.8) leads into a narrow tube (s.d), which gradually
dilates into a large sac (s). The narrow part is lined by small
epithelial cells, which are directly continuous with and perfectly
similar to those of the epidermis. The sac itself, which forms a
kind of bladder or collecting vesicle for the organ, is provided
with an extremely thin wall, lined with very large flattened cells.
The second section of the nephridium is formed by the coiled
tube, the epithelial lining of which varies slightly in the different
parts. The third section (s.0.¢), constitutes the most distinct
portion of the whole organ. Its walls are formed of columnar
cells almost filled by oval nuclei, which absorb colouring matters
with very great avidity, and thus render this part extremely
Fig. 11.—Nephridium from the
9th pair of legs of P. capensis.
o.s, External opening of seg-
mental organ; p.f, internal
opening of *nephridium into
the body cavity (lateral com-
partment) ; s, vesicle of seg- ©
mental organ; s.c.l, s.¢.2,
8.€.3, 8.¢.4, successive regions
of coiled portion of nephri-
dium; s.0.¢, third portion of
nephridium broken off at p.f
from theinternal vesicle, which
is not shown.
conspicuous. The nuclei are arranged in several rows. It ends
by opening into a vesicle (Fig. 14, D), the wall of which is so
delicate that it is destroyed when the nephridium is removed
from the body, and consequently is not shown in Fig. 11.
The fourth and fifth pairs are very considerably larger than
those behind, and are in other respects peculiar. The great mass
of each organ is placed behind the leg on which the external
- opening is placed, immediately outside one of the lateral nerve-
cords. The external opening, instead of being placed near the
base of the leg, is placed on the ventral side of the third ring
(counting from the outer end) of the thicker portion of the leg.
It leads into a portion which clearly corresponds with the collect-
ing vesicle of the hinder nephridia. This part is not, however,
dilated into a vesicle. The three pairs of nephridia in the three
foremost pairs of legs are rudimentary, consisting solely of a
vesicle and duct. The salivary glands are the modified nephridia
of the segment of the oral papillae.
VOL. V C
18 PERIPATUS
Generative Organs.
Ma.e.—tThe male organs (Fig. 12) consist of a pair of testes
(te), a pair of vesicles (v), vasa deferentia (v.d), and accessory
glandular tubules (f/). All the above parts le in the central
compartment of the body cavity. In P. capensis the accessory
glandular bodies or crural glands of the last (17th) pair of legs
are enlarged and prolonged into an elongated tube placed in the
lateral compartment of the body cavity (a.g).
The right vas deferens passes under both nerve-cords to join
Fic. 12.—Male generative organs of Peripatus capensis, viewed from the dorsal surface.
(After Balfour.) a.g, Enlarged crural glands of last pair of legs ; ¥'.16, 17, last pairs
of legs ; f, small accessory glandular tubes ; y, common duct into which the vasa
deferentia open ; te, testis; v, seminal vesicle ; v.c, nerve-cord ; v.d, vas deferens.
the left, and form the enlarged tube (p), which, passing beneath
the nerve-cord of its side, runs to the external orifice. The
enlarged terminal portion possesses thick muscular walls, and
possibly constitutes a spermatophore maker, as has been shown to
be the case in P. V. Zealandiae, by Moseley. In some specimens
a different arrangement obtains, in that the left vas deferens
passes under both nerve-cords to join the right.
FEMALE.—The ovaries consist of a pair of tubes closely ap-
plied together, and continued posteriorly into the oviducts. The
oviducts, after a short course, become dilated into the uteruses,
which join behind and open to the exterior by a median
a | GENERATIVE ORGANS AND DEVELOPMENT 19
7
opening. The ovaries always contain spermatozoa, some of which
project through the ovarian wall into the body cavity. Sperma-
tozoa are not found in the uterus and oviducts, and it appears
probable that they reach the ovary directly by boring through
the skin and traversing the body cavity In the neotropical
species there is a globular receptaculum seminis opening by two
short ducts close together into the oviduct, and there is a small
receptaculum ovorum with extremely thin walls opening into the
oviduct by a short duct just in front of the receptaculum seminis.
The epithelium of the latter structure is clothed with actively
moving cilia. In the New Zealand species there is a receptaculum
seminis with two ducts, but the receptacula ovorum have not
been seen.
There appear to be present in most, if not all, the legs some
accessory glandular structures opening just externally to the
nephridia. They are called the crural glands. |
DEVELOPMENT.
As stated at the outset, Peripatus is found in three” of the
great regions, viz. in Africa, in Australasia, and in South America
and the West Indies. It is a curious and remarkable fact that
although the species found in these various localities are really
closely similar, the principal differences relating to the structure
of the female generative organs and to the number of the legs,
they do differ in the most striking manner in the structure of
the ovum and in the early development. In all the Austral-
asian species the egg is large and heavily charged with food-
yolk, and is surrounded by a tough membrane. In the Cape
species the eggs are smaller, though still of considerable size ; the
yolk is much less developed, and the egg membrane is thinner
‘though dense. In the neotropical species the egg is minute
and almost entirely devoid of yolk. The unsegmented uterine
ovum of P. Novae-Zealandiae measures 1°5 mm. in length by *8 mm.
in breadth; that of P. capensis is 56 mm. in length; and that
of P. Trinidadensis ‘(04 mm. in diameter. In correspondence
with these differences in the ovum there are differences in the
early development, though the later stages are closely similar.
But unfortunately the development has only been fully worked
1 See Whitman, Journal of Morphology, vol. i. 2 See below, p. 24.
20 PERIPATUS | CHAP.
¢.
vut in one species, and to that species—P. capensis—the follow-
ing description refers. The ova are apparently fertilised in the
ovary, and they pass into the oviducts in April and May. In
May the brood of the preceding year are born, and the new ova,
which have meanwhile undergone cleavage, pass into the uterus.
There are ten to twenty ova in each uterus. The segmentation
is peculiar, and leads to the formation of a solid gastrula, consisting
of a cortex of ectoderm nuclei surrounding a central endodermal
mass, which consists of a much-vacuolated tissue with some
Fic. 13.—A series of embryos of P. capensis. The hind end of embryos B, C, D is
uppermost in the figures, the primitive streak is the white patch behind the blasto-
pore. (After Sedgwick.) A, Gastrula stage, ventral view, showing blastopore.
B, Older gastrula stage, ventral view, showing elongated blastopore and primitive
streak. ©, Ventral view of embryo with three pairs of mesoblastic somites, dumib-
bell-shaped blastopore and primitive streak. D, Ventral view of embryo, in which
the blastopore has completely closed in its middle portion, and given rise to two
openings, the embryonic mouth and anus. The anterior pair of somites have
moved to the front end of the body, and the primitive groove has appeared on the
primitive streak. E, Side view of embryo, in which the hind end of the body has.
begun to elongate in a spiral manner, and in which the appendages have begun.
At, antenna ; d, dorsal projection ; p.s, preoral somite. F, Ventral view of head of
embryo intermediate between E and G. The cerebral grooves are wide and shallow..
The lips have appeared, and have extended behind the openings of the salivary
glands, but have not yet joined in the middle line. At, antennae; c.g, cerebral
groove ; 7, jaws ; j.s, swelling at base of jaws ; LZ, lips ; M, mouth ; o7.p, oral papillae ;
0.8, opening of salivary gland. G, Side view of older embryo with the full number
of appendages, to show the position in which the embryos lie in the uterus,
irrecularly-shaped nuclei. The endoderm mass is exposed at one
point—the blastopore (gastrula mouth). The central vacuoles
of the endoderm now unite and form the enteron of the embryo,
and at the same time the embryo elongates into a markedly oval
form, and an opacity—the primitive streak—-appears at the hind
end of the blastopore (Fig. 13,B). This elongation of the embryo
is accompanied by an elongation of the blastopore, which soon
becomes dumb-bell shaped (Fig. 13, C). At the same time the
mesoblastic somites (embryonic segments of mesoderm) have made
—<— so
nd a alee eh
a a
ie DEVELOPMENT 21
‘their appearance in pairs at the hind end, and gradually travel for-
ward on each side of the blastopore to the front end, where the
somites of the anterior pair soon meet in front of the blastopore (Fig.
13,D). Meanwhile the narrow middle part of the blastopore has
closed by a fusion of its lips, so that the blastopore is represented
by two openings, the future mouth and anus. A primitive groove
makes its appearance behind the blastopore (Fig. 13, D). At
this stage the hind end of the body becomes curved ventrally
into a spiral (Fig. 13, E), and at the same time the appendages
appear as hollow processes of the body wall, a mesoblastic
somite being prolonged into each of them. The first to appear
are the antennae, into which the praeoral somites are prolonged.
The remainder appear from before backwards in regular order,
viz. jaw, oral papillae, legs 1-17. The full number of somites
and their appendages is not, however, completed until a later
stage. The nervous system is formed as an annular thickening
of ectoderm passing in front of the mouth and behind the anus,
and lying on each side of the blastopore along the lines of the
somites. The praeoral part of this thickening, which gives rise to
the cerebral ganglia, becomes pitted inwards on each side (Fig. 13,
F,¢g). These pits are eventually closed, and form the hollow
ventral appendages of the supra-pharyngeal ganglia of the adult
(Fig. 9, d). The lips are formed as folds of the side wall of the
body, extending from the praeoral lobes to just behind the jaw
(Fig. 13, F, Z). They enclose the jaws (j), mouth (Jf), and
opening of the salivary glands (0.s), and so give rise to the buccal
cavity. The embryo has now lost its spiral curvature, and
becomes completely doubled upon itself, the hind end being in
contact with the mouth (Fig. 13, G). It remains in this position
until birth. The just-born young are from 10-15 mm. in length
_ and have green antennae, but the rest of the body is either quite
_ white or of a reddish colour. This red colour differs from the
colour of the adult in being soluble in spirit.
The mesoblastic somites are paired sacs formed from the
anterior lateral portions of the primitive streak (Fig. 13, C).
As they are formed they become placed in pairs on each side of
the blastopore. The somites of the first pair eventually obtain a
position entirely in front of the blastopore (Fig. 13, D). They
form the somites of the praeoral lobes. The full complement of
somites is acquired at about the stage of Fig. 13, E. The relations
22 : PERIPATUS ; CHAP.
of the somites is shown in Fig. 14, A, which represents a transverse
section taken between the mouth and anus of an embryo of the
stage of Fig. 13, D. The history of these somites is an exceed-
Setly. interesting one, and may be described shortly as follows :—
They divide into two parts—a ventral part, which extends into
Fig. 14.—A series of diagrams of transverse sections through Peripatus embryos to
show the relations of the coelom at successive stages. (After Sedgwick.) A, Early
stage: 1, gut; 2, mesoblastic somite ; no trace of the vascular space; endoderm
and ectoderm in contact. B, Endoderm has separated from the dorsal and ventral
ectoderm. The somite is represented as having divided on the left side into a
dorsal and ventral portion: 1, gut; 2, somite ; 3, haemocoele. ©, The haemocoele
(3) has become divided up into a number of spaces, the arrangement of which is
unimportant. The dorsal part of each somite has travelled dorsalwards, and now
constitutes a small space (triangular in section) just dorsal to the gut. The ventral
portion (2’) has assumed a tubular character, and has acquired an external opening.
The internal vesicle is already indicated, and is shown in the diagram by the thinner
black line: 1, gut; 2’, nephridial part of coelom; 3, haemocoele ; 3’, part of
haemocoele which will form the heart—the part of the haemocoele on each side of
this will form the pericardium ; 4, nerve-cord. D represents the conditions at
the time of birth; numbers as in C, except 5, slime glands. The coelom is re-
presented as surrounded by a thick black line, except in the ee which forms the
internal vesicle of the uephridium,
the appendage, and a dorsal part (Fig. 14, B). The ventral part
acquires an opening to the exterior just outside the nerve-cord,
and becomes entirely transformed into a nephridium (Fig. 14,
D, 2’). The dorsal part shifts dorsalwards and diminishes rela-
tively in size (Fig. 14, C). Its fate differs in the different parts
I SPECIES : 23
of the body. In the anterior somites it dwindles and disappears,
but in the posterior part it unites with the dorsal divisions of
contiguous somites of the same side, and forms a tube—the
generative tube (Fig. 14, D, 2). The last section of this tube
retains its connexion with the ventral portion of the somite, and
so acquires an external opening, which is at first lateral, but soon
shifts to the middle line, and fuses with its fellow, to form the
single generative opening. The praeoral somite develops the
rudiment of a nephridium, but eventually entirely disappears.
The jaw somite also disappears; the oral papilla somite forms
ventrally the salivary glands, which are thus serially homologous
with nephridia. The perivisceral cavity of Peripatus is, as in all
Arthropoda, a haemocoele. Its various divisions develop as
a series of spaces between the ectoderm and endoderm, and
later in the mesoderm. The mesoderm seems to be formed
entirely from the proliferation of the cells of the mesoblastic
somites. It thus appears that in Peripatus the coelom does not
develop a perivisceral portion, but gives rise only to the renal
and reproductive organs.
APPENDIX!
Preripatus, Guilding
Soft-bodied vermiform animals, with one pair of ringed antennae, one
pair of jaws, one pair of oral papillae, and a varying number of claw-bearing
ambulatory legs. Dorsal surface arched and more darkly pigmented than
the flat ventral surface. Skin transversely ridged and beset by wart-like
spiniferous papillae. Mouth anterior, ventral; anus posterior, terminal.
Generative opening single, median, ventral, and posterior. One pair of
simple eyes. Brain large, with two ventral hollow appendages; ventral
cords widely divaricated, without distinct ganglia. Alimentary canal simple,
uncoiled. Segmentally arranged, paired nephridia are present. Body cavity
is continuous with the vascular system, and does not communicate with the
- paired nephridia. Heart tubular, with paired ostia. Respiration by means
of tracheae. Dioecious; males smaller and generally less numerous than
females. Generative glands tubular, continuous with the ducts. Viviparous.
Young born fully developed. They shun the light and live in damp places
beneath stones, leaves, and bark of rotten stumps. ~ They eject when irritated
a viscid fluid through openings at the apex of the oral papillae. Distribu-
1 Cf., in addition to the works quoted on pp. 3,4: A. Willey, ‘‘ Peripatus novae-
britanniae,” in Zoological Results, i., Cambridge, 1898 ; L. Bouvier, ‘‘ Cont. & l'histoire
des Péripates américains,” Ann. Soc. Entomol. de France, \xviii., 1899; W. F.
Purcell, ‘‘ Anatomy of Opisthopatus cinctipes,” Annals of the S. African Museum,
ii. 1900. R. Evans, Quart. J. Micr. Sci. xliv., 1901, pp. 473, 539.
24 PERIPATUS CHAP.
tion: South Africa (Cape Colony, Natal, and the Gaboon), New Zealand,
Australia and Tasmania, New Britain, South and Central America and the
West Indies, the Malay Peninsula [and Sumatra ?}.
The genus Peripatus, so far as adult conformation is concerned, is a very
homogeneous one. It is true, as was pointed out by Sedgwick, that the species
from the same part of the world resemble one another more closely than
they do species from other regions, but recent researches have shown that
the line between them cannot be so sharply drawn as was at first supposed,
and it is certainly not desirable in the present state of our knowledge to
divide them into generic or subgeneric groups, as has been done by some zoolo-
gists! The colour appears to be highly variable in species from all regions ;
it is perhaps more constant in the species from the Neotropical region than
in those from elsewhere. The number of legs tends to be variable whenever
it exceeds 19 pregenital pairs ; when the number is less than that, it is usually,
though not always, constant. More constant points of difference are the form
of the jaws, the position of the generative orifice, the presence of a recepta-_
culum seminis and a receptaculum ovorum, the arrangement of the primary
papillae on the distal end of the feet, and above all the early development.
South African Species.— With three spinous pads on the legs and feet,
with two primary papillae on the anterior side and one on the posterior
side ; outer jaw with one minor tooth at the base of the main tooth, inner
jaw with no interval between the large tooth and the series of small ones ;
last fully developed leg of the male with enlarged crural gland opening on a
large papilla placed on its ventral surface ; coxal organs? absent ; the nephri-
dial openings of the 4th and 5th pairs of legs are placed in the proximal
spinous pad. Genital opening subterminal, behind the last pair of fully devel-
oped legs; oviduct without receptacula seminis or receptacula ovorum; the ter-
minal unpaired portion of the vas deferens short. Ova of considerable size, but
with only a small quantity of yolk. The embryos in the uterus are all nearly of
the same age, except for a month or two before birth when two broods overlap.
The following species are aberrant in respect of these characters: Peri-
patus (Opisthopatus) cinctipes, Purcell (Cape Colony and Natal), presents a few
Australasian features ; there is a small receptaculum seminis on each oviduct,
some of the legs are provided with well-developed coxal organs, the feet
have one anterior, one posterior, and one dorsal papilla, and the successive
- difference in the ages of the embryos in the uterus, though nothing like that
found in the Neotropical species, is slightly greater than that found in other
investigated African species. Several pairs of legs in the middle region of
the body are provided with enlarged crural glands which open on a large
papilla. Male with four accessory glands, opening on each side of and behind
the genital aperture. P. thollont Bouvier, Equatorial West Africa (Gaboon)
shows some Neotropical features; there are 24 to 25 pairs of legs, the
1 The following genera or subgenera have been proposed: Peripatus for the
Neotropical species, Peripatoides for the Australasian, Peripatopsis and Opisthopatus
for the African, Paraperipatus for the New Britain species, and Loperipatus for the
Malay species.
2 Coxal organs are furrows on the ventral surface of some of the legs, with tumid
lips and lined by smooth non-tuberculate epithelium. It appears that they can be
everted.
I SPECIES ; 25
genital opening is between the penultimate legs, and though there are only
three spinous pads, the nephridial openings of the 4th and 5th legs are
proximal to the 3rd pad, coxal organs are present, and the jaws are on the
Neotropical type; the oviducts have receptacula seminis. The following
South African species may be mentioned: P. capensis Grube, with 17 (rarely
18) pairs of claw-bearing legs; P. balfourt Sedgw., with 18 (rarely 19) pairs ;
P. moseleyt Wood-M., with 20 to 24 pairs,
Australasian Species.—With 14, 15, or 16 pairs of claw-bearing ambu-
latory legs, with 3 spinous pads on the legs, and nephridial opening of the
4th and 5th legs on the proximal pad ; feet with one anterior, one posterior,
and one dorsal primary papilla; inner jaw without diastema, outer with or
without a minor tooth. Last leg of the male with or without a large white
papilla on its ventral surface for the opening of a gland; marked papillae
for the crural glands are sometimes present on other legs of the male; well-
developed coxal glands absent. Genital opening between the legs of the last
pair ; oviducts with receptacula seminis, without receptacula ovorum ; the ter-
minal portion of the vas deferens long and complicated ; the accessory male glands
open between the genital aperture and the anus, near the latter. Ova large
and heavily charged with yolk and provided with a stoutish shell. The uterus
appears to contain embryos of different ages. Specimens are recorded from
West Australia, Queensland, New South Wales, Victoria, and New Zealand.
The Australasian species are in some confusion. The number of claw-
bearing legs varies from 14 to 16 pairs, but the number most often found is
15. Whether the number varies in the same species is not clear. There
appears to be evidence that some species are occasionally or normally oviparous,
and in the supposed oviparous species the oviduct opens at the end of a papilla
called from its supposed function an ovipositor, but the oviparity has not yet
been certainly proved as a normal occurrence. Among the species described
may be mentioned P. leuckarti Sanger, P. insignis Dendy, P. oviparus Dendy,
P. viridimaculatus Dendy, P. novae-zealandiae Hutton, but it is by no means
certain that future research will maintain these. Mr. J. J. Fletcher indeed is of
opinion that the Australian forms are all varieties of one species, P. leuckartt.
Neotropical Species.— With 3 to 5 spinous pads on the legs, nephridial
opening of the 4th and 5th legs usually proximal to the third pad, and
feet either with two primary papillae on the anterior side and one on the
posterior, or with two on the anterior and two on the posterior; outer jaw
with small minor tooth or teeth at the base of the main tooth, inner jaw
with diastema. A variable number of posterior legs of the males anterior to
the genital opening with one or two large papillae carrying the openings
of the crural glands; well developed coxal organs present on most of the
legs. The primary papillae usually divided into two portions. Genital
opening between the legs of the penultimate pair; oviduct provided with
receptacula seminis and ovorum; unpaired part of vas deferens long and
complicated ; accessory organs of male opening at the sides of the anus. Ova
minute, with little food-yolk ; embryos in the uterus at very different stages
of development. The number of legs usually if not always variable in the same
species ; the usual number is 28 to 32 pairs, but in some species 40 to 43
pairs are found. The Neotropical species appear to fall into two groups:
(1) the so-called Andean species, viz., those which inhabit the high plateaux
or Pacific slope of the Andes; in these there are 4 (sometimes 5) pedal
26 PERIPATUS - CHAP. I
papillae, and the nephridial openings of the 4th and 5th legs are on the 3rd
pad ; and (2) the Caribbean species, viz. the remaining Neotropical species,
in which there are 3 papillae on the foot and the nephridial openings of
the 4th and 5th legs are between the 3rd and 4th pads. The Andean species
are P. eisenit Wh., P. tuberculatus Bouvier, P. lankestert Bouv., P. quitensis
Schm., P. corradi Cam., P. cameranot Bouv., and P. balzant Cam.
Of the remaining species, which are the majority, may be mentioned, P.
edwardsit Blanch., P. jamaicensis Gr. and Cock., P. trinidadensis Sedgw.,
—P. torquatus Ken., P. imthurmi Scl.
New Britain Peripatus.—With 22 to 24 pairs of claw-bearing legs,
with three spinous pads on the legs, and nephridial openings of legs 4 and 5
(sometimes of 6 also) on the proximal pad; feet with one. primary papilla on
the anterior, one on the posterior side, and one on the dorsal side (median
or submedian) ; outer jaw with a minor tooth, inner jaw without diastema ;
crural glands absent ; well-developed coxal organs absent. Genital opening
subterminal, behind the last pair of legs; oviduct with receptaculum seminis,
without receptaculum ovorum; unpaired part of vas deferens very short ;
accessory glands two, opening medianly and dorsally. Ova small, -1 mm. in
diameter, with little yolk; the embryos are provided with large trophic
vesicles (Willey). Embryos in the uterus of very different ages and probably
born all the year round.
But one species known, P. novae-britanniae Willey.
Sumatran! Peripatus.—Peripatus with 24 pairs of ambulatory legs, and
4 spinous pads on the legs. The primary papillae of the Neotropical character,
with conical bases. Generative opening between the legs of the penultimate
pair. Feet with only two papillae. Single species. P. sumatranus Sedgw.
Peripatus from the Malay Peninsula.2— With 23 to 25 pairs of claw-
bearing legs, 4 spinous pads on the legs, and nephridial openings of legs 4
and 5 in the middle of the proximal pad or on its proximal side; feet with 2
primary papillae, one anterior and one posterior ; outer jaw with 2, inner jaw »
with 2 or 3 minor teeth at base of main tooth separated by a diastema from
the row of small teeth; crural glands present in male only, in the two pairs
of legs preceding the generative opening; coxal organs present. Genital
opening between the penultimate legs; oviduct with receptacula seminis and
ovorum ; unpaired part of vas deferens long; male accessory glands two, —
opening medianly between the legs of the last pair. Ova large with much
yolk and thick membrane, like those of Australasian species; embryos with
slit-like blastopore, and of very different ages in the same uterus, probably
born all the year round. The species are P. weldoni Evans, P. horsta Evans,
and P. butlert Evans. It will thus be seen that the Malay species while
resembling the Neotropical species in the generative organs, differ from these
in many features of the legs and feet, in the important characters furnished
by the size and structure of the ovum, and by the early development.
1 The existence of this species is doubtful. The description of it was taken
from a singe specimen. The evidence that this specimen was found in Sumatra is
not conclusive.
2 I am indebted to Mr. R. Evans and the Editors of the Quart. J. Mier.
Sci. for permission to see proofs of Mr. Evans’ papers in vol. xliv. of that
journal.
‘MYRIAPODA: te |
RY be, ri :
F. G. SINCLAIR, M.A. — / he
(wormerty F, G. HEATHCOTE) |
‘Trinity College, Cambridge. ;
s .
¢ . 3 i oO
CHAPTER II
MYRIAPODA
INTRODUCTION —— HABITS—— CLASSIFICATION STRUCTURE—— CHILO-
GNATHA—CHILOPODA —— SCHIZOTARSIA —- SYMPHYLA— PAUR-
OPODA——EMBRYOLOGY——PALAEON TOLOGY.
TRACHEATA With separated head and numerous, fairly similar
segments. They have one pair of anterinae, two or three ‘pairs
of mouth appendages, and numerous pairs of legs.
The Myriapoda are a class of animals which are widely
distributed, and are represented in almost every part of the
globe. Heat and cold alike seem to offer favourable conditions
for their existence, and they flourish both in the most fertile
and the most barren countries.
They have not attracted much notice until comparatively
recent times. Compared with Insects they have been but little
known. The reason of this is not hard to find. The Myriapods
do not exercise so much direct influence on human affairs as
do some other classes of animals; for instance, Insects. They
include no species which is of direct use to man, like the silk-
worm or the cochineal insect, and they are of no use to him as
food. It is true that they are injurious to his crops. For instance,
the species of Millepede known as the “ wire worm” ! is extremely
harmful; but this has only attracted much notice in modern
times, when land is of more value than formerly, and agricul-
ture is pursued in a more scientific manner, and the constant
endeavour to get the utmost amount of crop from the soil has
caused a minute investigation into the various species of
animals which are noxious to the growing crop. The species of
1 Not to be confused with the larva of Elater lineatus, also known as “ wire-worm.”
30 MYRIAPODA’ CHAP. .
Myriapoda best known to the ancients were those which were
harmful to man on account of their poisonous bite.
Some writers have supposed that the word which is trans-
lated “mole” in the Bible (Lev. xi. 30) is really Scolopendra
(a genus of Centipede), and, if this is so, it is the earliest men-
tion of the Myriapods. They were rarely noticed in the classical
times; almost the only mention of them is by A®lian, who says
that the whole population of a town called Rhetium were driven
out by a swarm of Scolopendras. Pliny tells us of a marine Scolo-_
pendra, but this was most probably a species of marine worm.
Linnaeus included Myriapods among the Insects; and the
writers after him till the beginning of this century classed them
with all sorts of Insects, with Spiders, Scorpions, and even among
Serpents. It was Leach who first raised them to the importance
of a separate class, and Latreille first gave them the name
of Myriapoda, which they have retained ever since.
Myriapods are terrestrial animals, crawling or creeping on
the ground or on logs of wood, or even under the bark of trees.
There is, however, a partial exception to this; various naturalists
have from time to time given descriptions of marine Centipedes.
These are not found in the sea, but crawl about on the shore,
where they are submerged by each tide. A Geophilus of this
sort has been found in Jersey by Mr. Sinel,’ thus living a semi-
aquatic life. Professor F. Plateau, experimenting on the effect of
immersion on the Geophilidae, found that they could exist in sea
water from twelve to seventy hours, and in fresh water from six
to ten days. They thus offer a striking example of the power that _
their class possess of existing under unfavourable circumstances.
With regard to their habits the different species differ very
considerably. On the one hand we have the Chilopoda, or
Centipedes, as they are called in this country, active, swift, and
ferocious ; living for the most part in dark and obscure places,
beneath stones, logs of wood, and dried leaves, etc., and feeding
on living animals. On the other hand, we have the Chilognatha,
or Millepedes, distinguished by their slow movements and .
vegetable diet; inoffensive to man, except by the destruction
they occasion to his crops, and having as a means of defence no
formidable weapon like the large poison claws of the Centipedes,
but only a peculiarly offensive liquid secreted by special glands
1 See Nature, xli., 1890, p. 104.
II HABITS AND DISTRIBUTION 31
known by the unpleasant though expressive name of “stink glands,”
or by the more euphonious Latin name of glandulae odoriferae.
As a general rule the larger species of Myriapods are found
in the hotter climates, some of the tropical species being very
large, and some, among the family of the Scolopendridae, extremely
poisonous ; and it is even said that their bite is fatal to man.
If, however, the Centipede is sometimes fatal to man, it does
Fic. 15.—Scolopendra obscura. (From ©. L. Koch, Die Myriapoden.)
not always have it its own way, for we read of man making
food of Centipedes. It is hard to believe that any human being
could under any circumstances eat Centipedes, which have been
described by one naturalist as “a disgusting tribe loving the
darkness.” Nevertheless, Humboldt informs us that he has seen
the Indian children drag out of the earth Centipedes eighteen
inches long and more than half an inch wide and devour them.
Fic. 16.—Chordewma sylvestre. Lisp C. L. Koch, Die Myriapoden.)
This, I believe, is the only account of human beings using
the Myriapoda as food, if we except the accounts of the religious
fanatics among the African Arabs, who are said to devour Centi-
pedes alive; though this is not a case of eating for pleasure, for
the Scolopendras are devoured in company with leaves of the
prickly pear, broken glass, etc., as a test of the unpleasant things
which may be eaten under the influence of religious excitement.
32 MYRIAPODA CHAP.
A cold climate, however, is not fatal to some fairly large
species of Centipedes. A striking instance of this came under
my own observation some years ago. In 1886 I was travelling
in the island of Cyprus—the “ Enchanted Island,” as Mr. Mallock —
calls it in his book written about the same time—with the
intention of observing its natural history. This island consists
of a broad flat country crossed by two mountain ranges of con-
siderable height, thus offering the contrast of a hot climate in
the plains and a cold climate in the mountains. On the plain
country I found among the Myriapoda that the most common
species were a large Scolopendra and a large Lithobius. The
Scolopendra was fairly common, living for the most part under |
large stones, and it was a pleasant task to search for them in a
ruined garden near Larnaca.
This garden was made for the public, and is situated about a —
quarter of a mile from the old town of Larnaca. It has been
suffered to fall into decay, and is now quite neglected. Mr.
“Mallock has described many beautiful scenes in his book, but I
think he could have found few more beautiful than this old
garden with its deserted gardener’s house, now a heap of ruins,
but overgrown with masses of luxuriant vegetation, with beauti-
ful flowers peeping out here and there as if charitably endeavour-
ing to hide the negligence of man, and to turn the desolation
into a scene of beauty. I got several prizes in this garden, but
found the Myriapods were principally represented by the species
I have mentioned. |
After leaving Larnaca I rode across the plain country
through blazing heat, which was rapidly parching up the ground
to a uniform brown colour. At every stopping-place I found
the same species of Scolopendra and of Lithobius. After a few
days I began to get up among the mountains of the northern
range, and the burning heat of the treeless plain was gradually
exchanged for the cool shade of the pine-trees and the fresh air
of the mountains. As I ascended higher and higher the tem-
perature grew cooler till I reached the top of Mount Troodos, the
ancient Olympus. Here in the month of May the snow still
lingered in white patches, and the air was clear and cold. I
remained on the top of Troodos for a week, whilegl made a close
examination of the fauna to.be found fiere. I was much
surprised to find the identical species of Scolopendra and
u HABITS AND DISTRIBUTION 33
Inthobius with which I had become acquainted in the heat of
the low country, quite at home among the snow, and as common
as in, what I should have imagined to be, the more congenial
climate. Nor were they any the less lively. Far from exhibit-
_ ing any sort of torpor from the cold, the first one which I
_ triumphantly seized in my forceps wriggled himself loose and
_ fastened on my finger with a vigour which made me as anxious
to get rid of him as,I had formerly been to secure him. How-
_ ever, he eventually went into my collecting box.
On the whole, we may say that the Chilopoda are most
_ largely represented in the hotter climates, where they find a
- more abundant, diet in the rich insect life of the tropical and
_ semi-tropical countries. The more brightly-coloured Myriapods,
too, are for the most part inhabitants of the warmer countries.
The ease with which they are introduced into a country in the
earth round plants, and in boxes of fruit, may account to a great
extent for the wide distribution of the various species in
different countries. Mr. Pocock, who examined the Myriapods
brought back from the “ Challenger ” Expedition, informs us that
of ten species brought: from Bermuda, four had been introduced
from the West Indies. There is no doubt that animals which
can bear changes of temperature and deprivation of food, and
even a short immersion in the water, are well calculated to be
introduced into strange countries in many unexpected ways. |
As might be expected from a class of animals so widely
distributed, Myriapods show an almost infinite variety of size
and.colour. We find them so small that we can hardly see
them with the naked eye, as in the case of the tiny Polyxenus,
the Pauropidae, and the Scolopendrellidae. We also find them
more than six inches in length, as the larger species of Scolo-
pendridae. I am afraid we must dismiss as an exaggeration an
‘account of Centipedes in Carthagena a yard in length, and more
than six inches in breadth. The giver of this account— Ulloa
—informs us that the bite of this gigantic serpent-like creature
is mortal if a timely remedy be not applied. It is certainly
extremely probable that the bite of a Centipede of this size would
be fatal to any one. Some Centipedes are short and broad, and
composed of few segments, as Glomeris ; some are long and thin,
with more than a hundred segments, as Geophilus. They may be
beautifully coloured with brilliant streaks of colour, as in some
VOL. V D
34 MYRIAPODA CHAP.
of the Julidae or Polydesmidae, or may be of a dull nse rusty
iron colour, or quite black.
One of the strangest peculiarities found among Myriapadl is
that some of them (eg. Geophilus electricus) are phosphorescent.
As I was walking one summer evening near my home in
Cambridgeshire I saw what I thought was a match burning.
Looking more closely, I saw it move, and thinking it was a
glow-worm I picked it up, and was surprised to find that it was
a Geophilus shining with a brilliant phosphorescent light. I let
it crawl over my hand, and it left a bright trail of light behind
it, which lasted some time. I have been told that this species
is common in Epping Forest; also in Cambridgeshire.’
Besides G. electricus, G. phosphoreus has been described as
a luminous species by Linnaeus, on the authority of a Swedish
sea captain, who asserted that it dropped from the air, shining
like a glow-worm, upon his ship when he was sailing in the
Indian Ocean a hundred miles from land.
What the use of this phosphorescence may be is not Enotes with
any degree of certainty. It may be either a defence against
‘einen, or else a means of attracting the two sexes to one another.
The places which the Myriapods select for their habitation
vary as much as their colour and size, though, with a few excep-
tions, they chose dark and obscure places. A curious species of
Myriapod is Pseudotremia cavernarum (Cope), which is found in
certain caves in America. The peculiar life it leads in these
caves seems to have a great influence on its colour, and also
affects the development of its eyes. Mr. Packard’s account
of them is worth quoting: “Four specimens which I collected
in Little Wyandotte cave were exactly the same size as
those from Great Wyandotte cave. They were white tinged,
dusky on the head and fore part of the body. The eyes are
black and the eye-patch of the same size and shape, while the
antennae are the same.
“Six specimens from Bradford cave, Ind. (which is a small
grotto formed by a vertical fissure in the rock, and only 300 to
400 yards deep), showed more variation than those from the two
Wyandotte caves. They are of the same size and form, but
slightly longer and a little slenderer. ... The antennae are
much whiter than in those from the Wyandotte caves, and the
! See L. Jenyns’ Observations in Nat. Hist. London, 1846, p. 296.
II: HABITS AND DISTRIBUTION 35
head and body are paler, more bleached out than most of the
Wyandotte specimens... . It thus appears that the body is
most bleached and the eyes the most rudimentary in the Bradford
eave, the smallest and most accessible, and in which consequently
there is the most variation in surroundings, temperature, access of
light and changed condition of air. Under such circumstances
as these we should naturally expect the most variation.”
A strong contrast to these animals is afforded us by the
Scutigeridae (Schizotarsia). They are unknown in this country,
but abound in some of the Mediterranean countries and in parts
of Africa. They remind one strongly of spiders, with their long
Fic. 17.—Cermatia (Scutigera) variegata. (From C. L, Koch, Die Myriapoden. )
legs and their peculiar way of running on stones and about the
walls of houses.
Some years ago I was in Malta, and I used to go and watch
them on the slopes outside Valetta, where they were to be found
in great numbers. They used to come out from beneath great
‘stones and run about rapidly on the ground or on the stones and
rubbish with which the ground was covered, now and again
making a dart at some small insect which tempted them, and
seemingly not minding the blazing sun at all. As might be
expected from their habits, their eyes, far from being rudi-
mentary, like those of the cave-living Pseudotremia, or absent
1 « 4 Revision of the Lysiopetalidae, a family of the Chilognath Myriapoda, with
a notice of the genus Cambala,” by A. S. Packard, junior, Proc. Amer, Phil. Soc.
xxi. 1884, p. 187.
36 MYRIAPODA CHAP.
like those of the Polydesmidae, or of our own Cryptops, are
highly developed, and form the only example among the
Myriapods of what are known as facetted eyes. The Scutigeridae
are also remarkable among Myriapods for the possession of a
peculiar sense-organ which is found in no other Myriapod.
The Myriapods most numerous in our own country are
Lithobius and Julus. nthobius, which will be described later on,
may be found in almost any garden under dried leaves, stones,
etc. Julus, the common wire-worm, is found crawling on plants
and. leaves and under the bark of trees, and does a good deal of.
damage in a garden. Polydesmus is also frequently found in
great numbers, and usually a great many of them together.
Glomeris is also found, though it is not so common as the first
two mentioned animals. Geophilus is also common, and especially
in the south of England. Scolopendridae are only represented
by a single genus, Cryptops, which is not very common, though
by no means rare. The best place to find them is in manure
heaps. ‘The animals of this species are small compared to most
Scolopendras, and have the peculiarity of being without any eyes.
Scutigera is unrepresented in this country. One was found
in Scotland some years ago by Mr. Gibson Carmichael, but was
shown to have been imported, and not bred in the place.
The means of defence possessed by these animals also differ
very much in the different species of Myriapods. In the.
Centipedes the animals are provided with a powerful weapon in
the great poison claws which lie just beneath the mouth, and
which are provided with large poison glands, which supply a
fluid which runs through a canal in the hard substance of the
claw and passes into the wound made by the latter. The effect
of this fluid is instantaneous on the small animals which form
the food of the Centipedes. I have myself watched Lithobius in
this country creep up to a blue-bottle fly and seize it between the
poison claws. One powerful nip and the blue-bottle was dead, as
if struck by lightning. I have also seen them kill worms and also
other Lithobius in the same way. When another Lithobius is
wounded by the poison claws it seems to be paralysed behind
the wound. The Millepedes, on the other hand, have no such
offensive and defensive weapon. They rely for protection on the
fluid secreted by the stigmata repugnatoria (or glandulae odori-
jferae) mentioned before. This fluid has been shown to contain
oe HABITS, BREEDING 37
prussic acid, and has a very unpleasant odour. Most of the
Millepedes are provided with these glands; but in the cave
Myriapods mentioned before, the animals have not to contend
against so many adversaries, and these glands almost disappear.
Other Myriapods defend themselves by means of the Jong and
stiff bristles with which they are pro-
vided, eg. the little Polyxenus. This
means of defence seems to have been
more common among the fossil Myria-
pods than among those still living.
Variations in the shape and size of the
limbs are numerous,as might beexpected F1¢-18.—Polyxenus lagurus (From
: , C. L. Koch, Die Myriapoden).
in so large a class of animals. One of
the most curious of such variations is found in a Centipede of the
Scolopendra tribe, called Hucorybas, in which the last limbs are
flattened out and provided with paddle-shaped lobes. The use
of these is unknown, but it is probable that they are concerned
in some way with the breeding habits of the animal. The
habits of the Myriapods connected with their breeding are most
interesting, but have been very insufficiently investigated. There
is no doubt that a full inquiry into all such habits would be of
great interest, and would help to answer some of the problems
which are still unsolved in these forms. My own observations
refer to two forms—Julus terrestris among the Millepedes, and
Lithobius forficatus among the Centipedes. Julus terrestris is
one of the most common of the English Millepedes, and can be
easily obtained. I kept them in large shallow glass vessels with
a layer of earth at the bottom, and thus was able easily to
watch the whole process. They breed in the months of May,
June, and July. The female Ju/us when about to lay her eggs
. sets to work to form a kind of nest or receptacle for her eggs.
She burrows down into the earth, and at some distance below
the surface begins the work. She moistens small bits of earth
with the sticky fluid secreted by her salivary glands, which
become extraordinarily active in the spring. She works up
these bits of earth with her jaws and front legs till they are of
a convenient size and shape, and places them together. When
complete, the nest is shaped like a hollow sphere, the inside
being smooth and even, while the outside is rough and shows
the shape of the small knobs of earth of which it is composed.
38 MYRIAPODA CHAP.
She leaves a small opening in the top. The size of the whole
nest is about that of a small nut. When she is ready to lay her
eggs she passes them through the hole in the top, and usually
lays about 60 to 100 eggs at a time. The eggs, which are very
small, are coated with a glutinous fluid which causes them to
adhere together. When they are all laid she closes up the
aperture with a piece of earth moistened with her saliva; and
having thus hermetically sealed the nest, she leaves the whole to
its fate. The eggs hatch in about twelve days. |
A German naturalist, Dr. Verhoeff, has lately found that the
males of some Julidae undergo certain changes in the form of the
legs and other organs in autumn and spring. These changes are
probably connected with the breeding of the animal, and remind
us of the changes undergone during the breeding season by salmon
among the fishes.
Julus breed very readily if carefully attended to and well
supplied with food. If they cannot obtain the food they like
they will not breed so well. I found that sliced apples with
leaves and grass formed the best food for them.
The process in the case of Zithobius is much harder to watch.
Lithobius is not so plentiful as Julus terrestris, and the animals
are more impatient of captivity, more shy in their habits, and do
not breed so readily.
In January 1889 I was given the use of a room in the New
Museums at Cambridge, and was allowed to fit it up as I liked,
so that I was able to try the effect of different degrees of light
and darkness, and of different degrees of warmth. I succeeded
in observing the whole process. The female Zithobius is
furnished with two small movable hooks at the end of the
under surface of the body close to the opening of the oviduct.
These small hooks have been observed by many naturalists, but
their use has, so far as I know, never been described before.
They play an important part in the proceedings following the
laying of the egg. The time of breeding in Lithobius is rather
later than in Julus, and begins about June and continues till
August. There are first of all some convulsive movements of the
last segments of the body, and then in about ten minutes the
egg appears at the entrance of the oviduct. The egg is a
small sphere (about the size of a number five shot), rather
larger than that of Julus, and is covered with a sticky slime
o
. ieee HABITS, BREEDING 39
secreted by the large glands inside the body, usually called the
accessory glands. When the egg falls out it is received by
the little hooks, and is firmly clasped by them. This is
the critical moment in the existence of the JLithobius into
which the egg is destined to develop. If a male Lithobius sees
the egg he makes a rush at the female, seizes the egg, and at
once devours it. All the subsequent proceedings of the female
seem to be directed to the frustration of this act of cannibalism.
As soon as the egg is firmly clasped in the little hooks she
rushes off to a convenient place away from the male, and uses
her hooks to roll the egg round and round until it is completely
covered by earth, which sticks to it owing to the viscous material
with which it is coated; she also employs her hind legs, which
have glands on the thighs, to effect her purpose. When the
operation is complete the egg resembles a small round ball of
mud, and is indistinguishable from the surrounding soil. It is
thus safe from the voracious appetite of the male, and she leaves
it to its fate. The number of eggs laid is small when compared
with the number laid by. Julus.
The food in the case of ZLithobius consisted of worms and
blue-bottles, which were put alive into the glass vessel containing
the Lithobius. I tried raw meat chopped up, but they did not
thrive on it in the same way that they did on the living animals.
I also put into their vessel bits of rotten wood containing larvae
of insects, ete.
I have succeeded in bringing back some specimens of Polydesmus
alive from Madeira, and in getting them to breed in this country
—of course in artificial warmth—and their way of laying eggs
and making a nest resembles that of Julus. Geophilus has one
eurious habit in connexion with the fertilisation of the female.
The male spins a web and deposits in the middle of it a single
spermatophore, and the female comes to the web to be fertilised.
The Scolopendridae are said to bring forth their young alive, but
I think the evidence for this is unsatisfactory. What have
been taken for the young Scolopendrae are perhaps the large
spermatophores of the male, which are not unlike a larval Myria-
pod in size and shape. I have never been able to observe the
process of breeding in this family. I have had the spermatophores
sent me from Gibraltar as “eggs,” but a little examination soon
showed me their real character.
40 MYRIAPODA CHAP.
The mode of progression in the Myriapods differs considerably,
as might be expected in a class in which the number of legs
varies to such an extent. The swiftest among them are the
Scutigeridae with their long spider-like legs. The Scolopendridae
are also able to move with considerable rapidity, and are also
able to move tail forward almost as well as in the ordinary
manner. Where there are such a number of legs it becomes a
curious question as to the order in which the animal moves
them; and though several people have endeavoured to find this
out, the number of legs to be moved and the rapid movements
have rendered accurate observation impossible.
Some years ago Professor E. Ray Lankester tried to study the
order in which the legs of Centipedes moved, and came to the
conclusion (recorded in an amusing letter in Nature, 23rd May
1889) that if the animal had to study the question itself, it
would not get on at all. He finishes his letter with the follow-
ing verses :—
A Centipede was happy quite
Until a toad in fun
Said, ‘‘ Pray which leg moves after which ?”
This raised her doubts to such a pitch,
She fell exhausted in the ditch,
Not knowing how to run,
The progression of Millepedes is much slower than that of
the Centipedes, and it is remarkable that when the animal is in
motion a sort of wave runs down the long fringe-like row of feet.
I have endeavoured to make out this motion, but have never
been able to understand it satisfactorily. My belief was that
“the feet were moved in sets of five.
This wave-like peculiarity of motion is described in a curious
old book, An Essay towards a Natural History of Serpents.
Charles Owen, D.D. London, 1742: “The Ambua, so the
natives of Brazil call the Millepedes and the Centipedes, are
serpents. Those reptiles of thousand legs bend as they crawl
along, and are reckoned very poisonous. In these Multipedes the
mechanism of the body is very curious; in their going it is
observable that on each side of their bodies every leg has its
motion, one regularly after another, so that their legs, being
numerous, form a kind of undulation, and thereby communicate
to the body a swifter progression than one could imagine where
II NAMES FOR MYRIAPODS 4I
so many short feet are to take so many short steps, that follow
one another rolling on like the waves of the sea.”
Before proceeding to the classification of Myriapods, which
will form the next part of this account, a few words on the
common names for them may not be without interest.
In English we have the names Centipede and Millepede,
and the Continental nations have similar names implying the
possession of a hundred or a thousand legs, as the German
“ Tausendfiisse ” and the French “ Millepieds.” Of course these
are general words, simply implying the possession of a great
number of legs. But we have also among the peasantry a name
for Centipedes which conveys a much more accurate idea of the
number. The people of the eastern counties (I daresay the
term is more widely spread) call them “ forty legs.” This is not
quite accurate, but as Lithobius has 17 legs on each side, and
Scolopendra (Cryptops is the English species) has 21 on each side,
it is a better approximation than Centipede. But another
country has a still more accurate term. I found some Scolo-
pendra in Beyrout, and asked my native servant what he called
them. He gave them what I afterwards found was the common
Arab name for them, “‘arba wal ‘arbarin,” forty-four legs. Now
the Scolopendras, which in hotter climates are the chief representa-
-tives of the Centipedes, have actually forty-two legs, or, if the
poison claws are counted, forty-four. In looking up the Arab
term for Centipede I came across a curious description given of
_ them by Avicenna, the great Arabian physician: “This is an
animal known for its habit of going into ears. For the most part
it is a palm’s length” [about four inches, which is the average
length of many species]. “ On each side of the body it has twenty-
two feet, and moves equally well either backwards or forwards.”
With regard to its alleged habit of going into ears, the
learned Arabian has evidently made a false imputation on the
character of our animal, and has probably relied too much on the
stories told him. He has also exaggerated in stating that it
goes equally well either backwards or forwards. Some Centi-
pedes can go backwards very easily and well, though not so well
as forwards. Perhaps he preferred examining dead specimens,
which afford an easy opportunity of counting their legs, to experi-
menting with living animals, which might have resented liberties
taken with them.
42 MYRIAPODA CHAP,
The Persians have several words for them, less accurate than
the Arabs and more like our own terms. For instance, they call
them “Hazarpa,” or thousand feet, like our Millepedes; also
“Sadpa,” or hundred feet, equivalent to our Centipedes. Another
term resembles our common term before mentioned, “ Chehlpa,”
forty feet. A more figurative term is “tasbih dud,’ a worm
resembling a rosary with a hundred beads; this word is trans-
lated in Richardson’s Persian Dictionary as “a venomous insect
having eight feet and a piked tail.”
Classification of the Myriapoda.
Two of the principal writers on the classification of the
Myriapods are Koch and Latzel, both of whom have classified
the whole group. I do not wish for a moment to undervalue
the many authors who have done excellent work on the classifi-
cation of different groups and families, but I wish here to give
an outline of a classification of the whole class, and I naturally
have recourse to the authors who have treated the subject as a
whole. | |
Koch’s two works, the System der Myriapoden: and Die
Myriapoden, cover the whole range of the class, and his divisions
are clearly marked out and are easily understood, but both works
are comparatively old. He does not include the Scolopendrellidae
or the Pauropidae, which are now included by all naturalists in
the Myriapoda. Latzel is a more recent writer, and though his
work is entitled Zhe Myriapods of the Austro-Hungarian Empire,*
he gives much information about Myriapods not found in
Europe, and his work is fairly entitled to be considered as
embracing the whole class. He divides the Myriapods into four
Orders, including the Scolopendrellidae and Pauropidae. On the
whole, I think it will be better here to take the classification of
Koch, and to add to it the two Orders before mentioned, viz.
Symphyla containing one family the Scolopendrellidae, and Pauro-
poda with one family the Pauropidae.
The Orders are as follows :—
1 C. L. Koch, System der Myriapoden. Regensberg, 1847.
2 C. L. Koch, Die Myriapoden. Halle, 1863.
3 Latzel, Die Myriapoden der Cisterreichisch-Ungarischen Monarchie. Wien,
1880.
I CLASSIFICATION 43
Order I. CHILOGNATHA (= DIPLOPODA).
Antennae short, 6, 7, or 8 joints. Eyes congregate, simple,
or none. Body rings consisting of dorsal scute, two pleurae, 2
or 4 laminae pedigerae. Odoriferous glands mostly present.
Genital orifice in male and female placed between 2nd and 3rd
segment. In male, auxiliary copulatory organs in last segment
or on 7th, 7th and 8th, or 8th.
Suborder 1. PsrLapsocnatHa. Body having no auxiliary copulatory
organs or odoriferous glands.
Family 1. Polyxenidae. (Fig. 18, p. 37.)
Antennae 8 joints. Somites 11, last with bundle of setae. 13 pairs of legs,
Male with penis.
Family 2. Glomeridae.
12 tergites. 17 pairs of legs. Ocelli in single row.
Fic. 19.—Glomeris marginata. (From C. L. Koch, Die Myriapoden.)
Family 3. Zephroniidae.
13 tergites. 21 pairs of legs. Eyes crowded together in a cluster.
Fic. 20.—Sphaerotherium grossum. (From C. L. Koch, Die Myriapoden.)
Family 4. Julidae.
Body cylindrical. More than 30 body rings) Many eyes crowded
together in a cluster. Odoriferous glands always present.
Fic. 21.—Julus nemorensis. (From C. L. Koch, Die Myriapoden.)
44 MYRIAPODA CHAP.
Family 5. Blanjulidae.
Thin cylindrical body with more .than 30 body rings. Eyes either
absent or in a simple row beneath the edge of the forehead.
Latin
Fia. 22.—Blanjulus guttulatus. (From C. L. Koch, Die Myriapoden. )
Family 6. Chordewnidae,
Resemble the Polydesmidae (Fam. 7), but the head is longer and less rounded
in the forehead. The antennae are placed more at the side of the head.
Eyes small and numerous, ina cluster. Somites 30 or 32. ° (Fig. 16).
Family 7. Polydesmidae,
Body cylindrical, with a lobe or keel on the posterior part of the upper
surface of the body ring. Somites 19 or 20. No eyes.
Fic, 23.—Polydesmus collaris. (From C. L. Koch, Die Myriapoden.)
Suborder 2. ConopognatHa. Family 8. Polyzoniidae:
Head small, eyes few or none. Mouth-parts degenerate, adapted for
sucking. Pieural scutes free or coalesced. Laminae pedigerae free. Somites
30 to 108. Ist, and 2nd somites with one pair of feet. 3rd or Ist and
2nd apodous. Foramina repugnatoria present, Auxiliary copulatory organ
in 7th somite.
Order II. CHiLopopa (or SYNGNATHA).
Antennae with many joints, at least 14. Only one pair of
legs to each body ring. The genital opening on the last ring
of the body. Bases of the legs widely separate.
There are four families in this Order :—
II CLASSIFICATION
45
Family 1. Lithobicdae.
Body short and depressed. Eyes many or few; or a single ocellus on
each side. Antennae many joints but shorter than body. Number of
Fic. 25.—Lithobius erythrocephalus. (From C. L. Koch, Die Myriapoden.)
spiracles fewer than pairs of feet. Strong anal legs. Number of somites 15.
} Family 2. Scolopendridae.
- Body elongate. Ocelli on each side 4, 2, or none. Antennae 17 to 23
joints, much shorter than body. Spiracles fewer than pairs of feet. Anal
legs long and strong, number of legs 21 or 23. (Fig. 15, p. 31.)
Family 3. Notophilidae.
Body very long. Somites 100 to 170. No eyes, Maxillary palps very
Fic. 26.—Notophilus taeniatus. (From C. L. Koch, Die Myriapoden.)
thick. Compact or very short limbs.
Terminal point of last limb without
claw.
46 MYRIAPODA CHAP,
Family 4. Geophilidae.
Body very long; legs 13 to 173 pairs. No eyes. Antennae 14 joints,
Ba
Fia. 27.—Geophilus longicornis. (From C. L. Koch, Die Myriapoden.)
shorter than body. Spiracles fewer than pairs of legs. Anal pleurae coxi-
form.
Order III. ScuIzoTARSIA.
The tarsi of all the legs multiarticulate. The eyes facetted.
Peculiar sense organ beneath the head.
Family 1. Scutigeridae.
Body short and strong. Antennae very long and thin. Facetted eyes.
No spiracles, but stomata in back. (Fig. 17, p. 35.)
Order IV. SYMPHYLA.
Body small, 12 segments, which according to Schmidt equal
22 primary segments. One pair of tracheae opening in the head.
Genital opening before coxae of 4th pair. 1st and 2nd segments
with one pair of legs, rest with a pair and parapodia. Ovaries
beneath the gut. A head artery and a dorsal vessel with ostia
and alary muscles.
Family 1. Scovopendrellidae.
With the characters of the Order.
" STRUCTURE 47
Order V. PAUROPODA:
Body 12 somites, 8 of which fuse to double somites, 7 pairs of
legs. 1st and anal legs with 5 joints, rest 6 joints. Antennae
branched. No tracheae or vascular system. Ovary below gut,
testis above. Genital opening in 3rd somite.
Family 1. Pawropidae.
With long legs.
Family 2. Eurypawropidae.
With short legs.
The Structure of the Myriapoda.
Having now given a short view of the classification of the
Class, I will proceed to give a general account of their structure,
the variations in which have led to the divisions into the various
Orders and Families. Their structure shows resemblances to
several widely different classes of animals. One cannot help
being impressed with their likeness to the Worms, at the same
time they have affinities with the Crustaceans, and still more
with the Insects. In the latter class the likeness of the Thy-
sanuridae to Scolopendrella and Pauropus have induced a cele-
_ brated Italian anatomist, Professor Grassi, to claim the former as
the ancestors of the Myriapoda.
Myriapods have a body which is segmented, as it is termed ;
that is, composed of a number of more or less similar parts or
segments joined together.
One of the most important characteristics which distinguish
Myriapods from other Arthropoda is the fact that they possess
on the posterior segments of the body true legs which are
jointed and take part in locomotion. The head is in all cases
quite distinct from the body, and may be regarded as a number
of segments fused together into one mass. Their heads are
always provided with a single pair of antennae and mouth
appendages, consisting of an upper lip, a pair of mandibles or
jaws, and one to two pairs of maxillae. The mandibles resemble
those of Insects, and are strongly toothed. In the Chilognatha
a pair of maxillae are fused so as to form a single oval appendage.
In the Chilopoda they each consist of a single blade bearing a
48 MYRIAPODA CHAP.
short palp or feeler. The mouth parts may have the forms
known as chewing, biting, or suctorial (Polyzonium) mouth
appendages. ;
With the exception of the terminal segment, and in many
cases the first or the seventh, each segment bears one or two
pairs of limbs. These may be very idug, as in Seutigera, or very
short, as in Polyxenus. They may be attached close to one
another near the ventral middle line of the body, or may
have their bases far apart from each other, as in the Chilopoda.
The exoskeleton or external armour is composed of chitin
(Chilopoda) or of chitin with calcareous salts deposited in it
(Chilognatha).
Their internal structure has a great likeness to that of
Insects.
The general position of the internal organs may be seen from ~
Fig. 28, which shows a Lithobius dissected so as to exhibit the
digestive and nervous systems.
The digestive canal, which is a straight tube, extends through-
out the whole length of the body, and terminates in the last
segment of the body. It may be divided into the ee
parts :—
1. A narrow oesophagus, beginning with the mouth or bisael
cavity, and receiving the contents of two or more
salivary glands (d).
2. A wide mesenteron or mid-gut (x) extending throughout
almost the whole length of the body.
3. A rectum which at its junction with the mid-gut receives
the contents of two or four Malpighian tubes (g, h) which
function as kidneys. Their function was for a long
" time unknown, but the discovery of crystals of uric
acid in them placed the matter beyond doubt.
The heart has the form of a long pulsating dorsal vessel
which extends through the whole length of the animal. It is”
divided into a number of chambers, which are attached to the
dorsal wall of the body, and are furnished with muscles of a
wing-like shape, which are known as the alary muscles, and
which govern its pulsations. The chambers are furnished with
valves and arteries for the exit of the blood, and slits known as
ostia for the return of the blood to the heart. The blood enters .
the chambers of the heart from the body cavity through the
1 STRUCTURE 49
ostia, and passes out through the arteries to circulate through
' the organs of the body and to return by the ostia.
The two figures below (Figs. 29 and 30) show the position of
the arteries and the ostia in a single segment of the body. The
heart is too small and delicate to be seen with the naked eye; it
es
Fic. 28.—Lithobius dis-
sected. (After Vogt
and Yung.) .
a, antennae.
6, poison claws.
c, brain.
d, salivary glands.
é, legs.
J, nerve cord.
g, Malpighian tube.
h, Malpighian tube.
7, vesicula seminalis.
js accessory gland,
k, accessory gland,
Z, testis. «
m, thigh gland.
n, digestive tube.
therefore requires the aid of the microscope. 7 A =< y ip
SOsScores
RY
ie Py A
Fic. 87.—Later stage: nw,
nucleolus ; ¢.p, nucleus ;
y.sp, yolk spherules ; ch,
shell.
cells, cannot divide like them, and therefore the segmentation of
the ovum in Myriapods is irregular, as it is called.
I will now go back a little and describe what happens to the
ovum before the process of segmentation iscomplete. It increases
in size and forms the supply of food yolk which is to provide the
uutriment of the ovum. Then after impregnation the egg-shell
is formed round it, and it becomes what we know as the egg.
This egg is not a perfect sphere, but is oval (in most Myriapods)
in shape. The egg is laid, and the process of segmentation begins
shortly after it is laid, as has already been described.
When it has been laid for about 36 hours, if we take an egg
and, after proper preparation, cut it into thin slices known to
FORMATION OF THE EGG: 65
scopists by the name of sections, and examine it by means
f the microscope, we shall see that segmentation, has resulted in
: is. Just beneath the egg-shell there is a thin layer of cells,
ne cell thick, which completely surrounds the egg. Inside
is coat of cells is the food yolk, with a few cells scattered
bo in it at rare intervals, something like the raisins in a
um-pudding.
"With the next process the formation of the young Myriapod
nay be said to begin. A strip along the’ length of the oval-
shaped egg is thickened, and this thick mass of cells represents
the future ventral surface of the animal. The rest of the thin
ee er of cells: already mentioned just below the shell will form
the shell or exoskeleton of the future animal. The thick strip
of cells at the ventral surface has by this time split into
layers, so that, resorting to our microscope again, a section through
“the short axis of the oval-shaped. egg—a transverse section—
- will show us— |
1. The egg-shell.
2. A layer of cells completely surrounding the egg, thin
everywhere but on the ventral surface. This layer is
known to embryologists as the epiblast. The thick
part of the epiblast on the venttat surface gives rise to
the nervous system.
3 and 4. Two layers of cells connected in the middle, along
the line of the thick strip, but separate elsewhere, and
a ~ not extending round the whole of the inside. These
; layers constitute what is: known asthe mesoblast, and
give rise to the muscles and most of ‘the internal
organs.
5. The scattered cells in the yolk, They’ at are novi as the
hypoblast and give rise to the digestive canal. 5
After this point is reached the formation of the organs
Been. The segments are formed in order from before back-
wards. First the head, then the next segment, and so on.
4 When the number of segments with which the animal will be
hatched are formed, another process begins, and the tail end of
= - he animal, which can already be distinguished, is bent towards
the head. This is a process that takes place in many animals
besides Myriapods, and is called the formation of the ventral
_ flexure. Shortly after this the animal bursts the shell and comes
VOL. V F
.
66 MYRIAPODA CHAP.
into the outer world. The various processes may be understood
by reference to the Figs. 36, 37, 38, 39, which are succes-
sive stages in the development of a Chilognath. Figs. 37, 38,
are thin slices through the shorter diameter of the egg, which, as
Fic. 38. — Transverse section
through next stage: mk,
keel-like mass of cells from
which the mesoblast is pro-
duced ; ec, epiblast. (From
Heathcote, Post. Emb. Dev.
of Julus terrestris; Phil.
Trans, vol. 179, 1888, B.)
before mentioned, is an oval in shape. Fig. 39 is a section
through the longer diameter of an egg in a more advanced stage
of development, in fact just about to burst the shell. The body
of the future animal is marked by constrictions, the future
segments. Some of the organs are already formed, as the brain
Fic. 39. — Longitu-
dinal section
through later stage:
Segs. 2, 3, etc., seg-
ments ; Ceph. Seg,
head; mes, meso-
blast ;en,hypoblast;
Q)
=e st,futuremouth; pr,
future anus ; mesen,
50° 6, gut; mem.ex, as in
® eee en. Fig. 41, (From
& Oy } eS r) Heathcote, Post.
Beceeies mesen. Emb. Dev. of Julus
Loe® eee? terrestris. )
and the digestive tube, the openings of which will form the
mouth (st) and the anus (pr).
Myriapods are hatched at different stages of development.
The Chilognatha have only three appendages, which are so
little developed that they are only small shapeless stumps, while
u FORMATION OF THE YOUNG ANIMAL 67
the Chilopoda have the full number of legs in some cases; in
others only a small number of legs, but yet more than the three
pairs of legs of the Chilognatha, and fully developed instead of
stump-like. The eyes are usually developed late in the life of
the young animal. The bursting of the egg-shell is assisted in
some Myriapods by a special kind of spike on the back part of
the head.
The Fig. 40 shows a young Chilognath which has just burst
the shell and come into the outer world. ya
Tt is still surrounded with a membrane a
which has been formed by its skin or
epiblast within the egg. One eye-spot has
been formed.
Fig. 41 shows a longitudinal section
through the young Chilognath shown in 2,
Fig. 40, and the next (Fig. 42) a transverse Fic. 40.—Young Julus ter-
section through the same. In comparing sg ay ccc
the two Figs. 41 and 42 it must be remembered that they are
ir yy
« ii Yili
a
Oe
Xx) :
“< ,
Ss
Fic. 47—Diagram of exterior of insect: the two vertical dotted lines indicate the
divisions between H, head; T, thorax; and A, abdomen: a, antenna; 0, labrum ;
c, mandible ; d, maxillary palpus ; e, labial palpus ; /, facetted eye ; g, pronotum ;
h, mesonotum ; 7, metanotum ; &, wings; J, to /,), abdominal segments ; m, the
internal membranous portions uniting the apparently separated segments ; m, cerci ;
o, stigma; py, abdominal pleuron bearing small stigmata; 9g), go, g3, pro-, meso-,
meta-sterna ; 7,, mesothoracic episternum ; s}, epimeron, these two forming the
mesopleuron ; 7, S,, metathoracic episternum and epimeron; ¢, coxa; v, trochanter ;
w, femur; x, tibia; y, tarsus ; z, gula.
the nature of an appendage, and the theory of a triple origin for
these segments may be dismissed. There are, however, several facts
that indicate a duplicity in these somites, among which we may
specially mention the remarkable constancy of two pleural pieces
on each side of each thoracic segment.. The hypothesis of these
rings being each the representative of two segments cannot there-
fore be at present considered entirely untenable, and in that case
the maximum and minimum numbers that can be suggested
appear to be twenty-four and eleven, distributed as follows :—
1 Stettin. Ent. Zeit. 1, 1889, p. 165.
STRUCTURE 89
Maximum. Minimum.
Head F ‘ 4, 3
Thorax. / Pre 3
Abdomen . . wll 5
_ Total . 24 11
} ealthdugh it is not ‘ecebabts that ultimately so great a difference
/ of these figures indicate will be found to prevail, it is certainly
ab present premature to say that all Insects are made up of the
| same number of primary segments.
_ A brief account of the structure of the integument will be
found in the chapter dealing with the post-embryonic develop-
_ ment.
_ The three great regions of the Insect body are functionally as
_ well as anatomically distinct. The head bears the most important
a Bor the sense organs, viz. the antennae and ocular organs ; it includes
a a greater of the nerve-centres, and carries the mouth as well
_ as the appendages, the trophi, connected therewith. The thorax
is chietly devoted to the organs of locomotion, bearing externally
___ the wings and legs, and including considerable masses of muscles,
: 4 as well as the nerve centres by which they are innervated ; through
_ the thorax there pass, however, in the longitudinal direction,
~ those structures by which the unity of the organisation is com-
cto, viz. the alimentary canal, the dorsal vessel or “ heart” for
i D distributing the nutritive fluid, and also the nerve cords. The
_ abdomen includes the greater part of the organs for carrying on
a the life of the individual and of the species; it also frequently
bears externally, at or near its termination, appendages that are
doubtless usually organs of sense of a tactile nature.
Tn the lower forms of Insect life there is little or no actual in-
~ ternal triple division of the body; but in the higher forms such
_ separation becomes wonderfully complete, so that the head may
communicate with the thorax only by a narrow isthmus, and the
thorax with the abdomen only by a very slender link. This .
x ee eement is carried to its greatest extreme in the Hymenoptera
- Aculeata. It may be looked on as possibly a means for separating
the nutrition of the parts included in the three great body
_ divisions.
4 Along each side of the body extends a series of orifices for the
admission of air, the stigmata or spiracles; there are none of
these on the head, but on each side of most of the other segments
2
i
os
4
90 , SEGMENTS CHAP.
there is one of these spiracles. This, however, is a rule subject
to many exceptions, and it is doubtful whether there is ever a
spiracle on the last abdominal segment. Even in the young stage
of the Insect the number of these stigmata is variable; while
in the perfect Insect the positions of some of the stigmata may be
much modified correlatively with the unequal development or
consolidation of parts, especially of the thorax when it is highly
modified for bearing the wings. |
The segments of the Insect are not separate parts connected
with one another by joints and ligaments; the condition of the
Insect crust is in fact that of a continuous long sac, in which
there are slight constrictions giving rise to the segments, the
interior of the sac being always traversed from end to end by a
tube, or rather by the invaginated ends of the sac itself which
connect with an included second sac, the stomach. The more
prominent or exposed parts of the external sac are more or less
hard, while the constricted parts remain delicate, and thus the
continuous bag comes to consist of a series of more or less hard
rings connected by more delicate membranes. This condition is
Fig. 48—Tillus elongatus, fully distended larva.
readily seen in distended larvae, and is shown by our figure 48
which is taken from the same specimen, whose portrait, drawn
during life, will be given when we come to the Coleoptera, family |
Cleridae. The nature of the concealed connexions between the
apparently separate segments of Insects is shown at m, Fig. 47,
p. 88.
As the number of segments in the adult Insect corresp6nds— ~
except in the head—-with the number of divisions that appear
very early in the embryo, we conclude that the segmentation of
the adult is, even in Insects which change their form very greatly
during growth, due to the condition that existed in the embryo ;
but it must not be forgotten that important secondary changes
occur in the somites during the growth and development of
the individual. Hence in some cases there appear to be more
than the usual number of segments, e.g. Cardiophorus larva,
and in others the number of somites is diminished by amal-
STRUCTURE 91
— or by the extreme reduction in size of some of the
‘Besides the division of the ecaee into consecutive segments,
4 pethict feature is usually conspicuous; the upper part, in many
‘a segments, being differentiated from the lower and the two being
. — connected together by intervening parts in somewhat the same sort
. of way as the. segments themselves are connected. Such.a differen-
_ tiation is never visible on the head, but may frequently be seen in
_ the thorax, and almost always in the abdomen. A dorsal and a
) _ ventral aspect are thus separated, while the connecting bond on
4 either side forms a pleuron. By this differentiation a second form
p of symmetry is introduced, for whereas there is but one upper and
_ one lower aspect, and the two do not correspond, there are two
x lateral and similar areas. This bilateral symmetry is conspicuous
in nearly all the external parts of the body, and extends to most
. of the internal organs. The pleura, or lateral regions of the
_ sac, frequently remain membranous when the dorsal and ventral
aspects are hard. The dorsal parts of the Insect’s rings are
a :. “also called by writers terga, or nota, and the ventral parts
4 -sterna..
___ ‘The appendages of the body are —(1) a pair of antennae ; (2)
the trophi, constituted by three pairs of mouth-parts; (3) three
pairs of legs; (4) the wings'; (5) abdominal appendages of various
a Kinds, but usually jointed. Before considering these in detail we
shall do well to make ourselves more fully acquainted with the
: Be piementary details of the structure of the trunk.
In the adult Insect the integument or crust of the body is
_ more or less hard or shell-like, sometimes, indeed, very hard, and
on ‘examination it will be seen that besides the divisions into
‘segments and into dorsal, ventral, and pleural regions, there are
=a a indicating the existence of other divisions, and it will be
found that by dissection along these lines distinct pieces can be
readily separated. Each hard piece that can be so separated is
- called a sclerite, and the individual sclerites of a segment have
received names from entomotomists. The sclerites are not really
o- '* The wings, by many morphologists, are not included in the category of
_ “appendages”; they apparently, however, differ but little in their nature from
_ legs, both being outgrowths of the integument; the wings are, however, always
___ post-embryonic in actual appearance, even when their rudiments can be detected in
_ the larva. No insect is hatched from the egg in the wing-bearing form.
92 HEAD CHAP.
quite separate pieces, though we are in the habit of speaking of
them as if such were the case. If an Insect be distended by
pressure from the interior, many of the sclerites can be forced
apart, and it is then seen that they are connected by delicate
membrane. ‘The structure is thus made up of hard parts meeting
one another along certain lines of union—sutures—so that the
original membranous continuity may be quite concealed. In
many Insects, or in parts of them, the sclerites do not come into
apposition by sutures, and are thus, as it were, islands of hard
matter surrounded by membrane. . A brief consideration of some
of the more important sclerites is all that is necessary for our
present purpose: we will begin with the head.
The head is most variable in size and form; as a part of its
surface is occupied by the eyes and as these organs differ in
shape, extent, and position to a surprising degree, it is not a
matter for astonishment that it is almost impossible to agree as
to terms for the areas of the head. Of the sclerites of the head
itself there are only three that are sufficiently constant and
definite to be worthy of description here. These are the clypeus,
the epicranium, and tlie gula. The clypeus is situate on the
upper surface of the head-capsule, in front; it bears the labrum
which may be briefly described as a sort of flap forming an upper
lip. The labrum is usually possessed of some amount of mobility.
The clypeus itself is excessively variable in size and form, and
sometimes cannot be delimited owing to the obliteration of the
suture of connexion with the more posterior part of the head;
it is rarely or never a paired piece. Occasionally there is a
more or less distinct piece
interposed between the
clypeus and the labrum,
and which is the source
of considerable difficulty, as
it may be taken for the
i clypeus. Some authors call
Fre. 49.—Capsule of head of beetle, Harpalus the clypeus the epitie but
caliginosus : A, upper ; B, under surface: a, it is better to use this latter
dypenss 2 gpicraninn: «, pofoeaniim ; term for the purpose of indi
men’; g, submentum ; h, cavity for insertion cating the part that is imme-
eh ast diately behind the labrum,
whether that part be the clypeus, or some other sclerite; the
‘inal i + -s
z oS. HEAD 93
term is very convenient in those cases where the structure cannot
Be, or has not been, satisfactorily determined morphologically.
= tn Figure 50 the parts usually visible on the anterior
aspect of the head and its appendages are shown so far as these
latter can be seen when the mouth is closed ; in the case of the
Insect here represented the bases of the mandibles are ecreatly
~~ seen (gy), while their apical portions are
entirely covered by the labrum, just
below the lower margin of which the
tips of the maxillae are seen, looking
as if they were the continuations of the
Be riandiblen
a The labrum is a somewhat perplex-
. ing piece, morphologists being not yet
agreed as to its nature; it is usually
_ placed quite on the front of the head,
and varies extremely in form; it is es
3 nearly always a single or unpaired hee aya: me leader 3
Be ‘piece ; the French morphologist Chatin epicranium; 6, compound
considers that it is really a paired Sake Pisce Ligh nats
structure. . g, base of mandible ; 4, max-_
ee the gula (Fig. 49,B d,and Fig 47, tis) P'sbesof manila.
_ #) is a piece existing in the middle
longitudinally of the under-surface of the head; in front it
pears the mentum or the submentum, and extends backwards
to the great occipital foramen, but in some Insects the gula
is in front very distant from the edge of the buccal cavity.
_ The epicranium forms the larger part of the head, and is con-
sequently most inconstant in size and shape ; it usually occupies
the larger part of the upper - surface, and is reflected to the
-_under-surface to meet the gula. Sometimes a transverse line
exists (F ig. 49, A) dividing the epicranium into two parts, the
_ posterior of which has been called the protocranium; which,
however, is nota good term. The epicranium bears the antennae ;
these organs do not come out between the epicranium and the
clypeus, the foramen for their insertion being seated entirely in
_ the epicranium (see Fig. 50). In some Insects there are traces of
5 the epicranium being divided longitudinally along the middle
line. When this part is much modified the antennae may
appear to be inserted on the lateral portions of the head, or even
94 INSECTS CHAP.
on its under-side; this arises from extension of some part of the
epicranium, as shown in Fig. 49, B, where h, the cavity of
insertion of the antenna, appears to be situate on the under-
surface of the epicranium, the appearance pee due to an
infolding of an angle of the part.
There is always a gap in the back of the head for the passage
of the alimentary canal and other organs into the thorax; this
opening is called the occipital foramen. Various terms, such as
frons, vertex, occiput, temples, and cheeks, have been used for
designating areas of the head. The only one of these which is of
importance is the gena, and even this can only be defined as the
anterior part of the lateral portion of the head-capsule. An
extended study of the comparative anatomy of the head-capsule
is still a desideratum in entomology. The appendages of the head
that are engaged in the operations of feeding are frequently
spoken of collectively as the trophi, a term which includes the
labrum as well as the true buccal appendages.
The appendages forming the parts of the mouth are paired,
and consist of the mandibles, the maxillae, and the labium, the
pair in this latter part being combined to form a single body.
The buccal appendages are frequently spoken of as gnathites.
The gnathites are some, if not all, of them composed of apparently
numerous parts, some of these being distinct sclerites, others
membranous structures which may be either bare or pubescent—
that is, covered with delicate short hair. In Insects the mouth
functions in two quite different ways, by biting or by sucking.
The Insects that bite are called Mandibulata, and those that suck
Haustellata. In the mandibulate Insects the composition of the
gnathites is readily comprehensible, so that in nearly the whole
of the vast number of species of that type the corresponding ©
parts can be recognised with something like certainty. This,
however, is not the case with the sucking Insects ; in them the
parts of the mouth are very different indeed, so that in some
cases morphologists are not agreed as to what parts really
correspond with some of the structures of the Mandibulata. At
present it will be sufficient for us to consider only the mandibulate
mouth, leaving the various forms of sucking mouth to be~
discussed when we treat of the Orders of Haustellata in detail.
The upper or anterior pair of gnathites is the mandibles,
(Fig. 50, 9). There is no part of the body that varies more than
fa
mm | MOUTH-PARTS _ oe
does the mandible, even in the mandibulate Insects. It can
searcely be detected in some, while in others, as in the male stag-
beetle, it may attain the length of the whole of the rest of the
body; its form, too, varies as much as its size ; most usually,
however, the pair of mandibles are somewhat of the form of
eallipers, and are used for biting, cutting, holding, or crushing
_ purposes. The mandibles are frequently armed with processes
spoken of as teeth, but which must not be in any way confounded
with the teeth of Vertebrates. The only Insects that. possess an
~ articulated tooth are the Passalidae, beetles armed with a rather
large mandible bearing a single mobile tooth among others that
are not so. Wood Mason and Chatin consider the mandibles to
Fic. 51. — Mandibles,
maxillae, and labium
of Locusta viridis-
sima; A, mandibles ;
B, maxillae (lateral
parts) and Jlabium
(middle parts) united :
a, cardo; b, stipes ;
c, palpiger ; d, max.
palp. ; ¢, lacinia; /,
galea; g,submentum ;
h, mentum; 7%, pal-
piger ; %, labial pal-
pus; 7, ligula; m,
paraglossa (galea); ,
lacinia ; 0, lingua.
be, morphologically, jomted appendages, and the latter authority
states that in the mandible of Embia he has been able to distin-
guish the same elements as exist in the maxillae. In aculeate
Hymenoptera the mandibles are used to a considerable extent
for industrial purposes.
The maxilla is a complex organ consisting of numerous pieces,
viz. cardo, stipes, palpiger, galea, lacinia, palpus. The galea and
lacinia are frequently cailed the lobes of the maxilla. The
maxilla no doubt acts as a sense organ as well as a ‘mechanical
apparatus for holding; this latter function being subordinate to
the other. In Fig. 68, p. 122, we have represented a complex
maxillary sense-organ.
The labium or lower lip has as its basal portion the un-
96 INSECTS | CHAP.
divided mentum, and closes the mouth beneath or behind,
according as the position of the head varies. In most Insects
the labium appears very different from the maxilla, but in many
cases several of the parts corresponding to those of the maxilla
can be clearly traced in the labium.
The mentum is an undivided, frequently very hard, piece,
continuous with either the submentum or the gula, and anterior
to this are placed the other parts, viz.
the labial palpi and their supports, the
palpigers; beyond and between these
exists a central piece (Fig. 52, B, e),
about whose name some difference of
opinion prevails, but which may be called
the ligula (languette of French authors),
and on each side of this is a paraglossa.
In the Orthoptera the single median
piece —the ligula of Coleopterists — is
represented by two divided parts. In
some Insects (many Coleoptera) there is
interposed between the mentum and the
palpigers a piece called the hypoglottis
(Fig. 52, B, 6). It is not so well ascer-
tained as it should be, that the pieces of
the lower lip bearing the same names in
different Orders are in all cases really
homologous, and comparison suggests that -
Fig. 52.—Maxilla and lower the hypoglottis of Coleoptera may pos-
lip of Coleoptera. A, Max- . " : ‘
illa of Passalus: a, cardo: Sibly represent the piece corresponding
saan ; idee ps ae to the mentum of Orthopterists, the so-
rior lobe or lacinia; f, Called mentum of beetles being in that
outer or superior lobe or ease the submentum of Orthopterists.
galea: B, Labium of Har- : &
palus caliginosus: a, men- There is another part of the mouth
bam 5 0 hyposlottis; © to which we may call special atten-
palpiger (support of the : .
labial palp); d, palp; e, tion, as it has recently attracted more
Aigule 5. fy: Dante hones attention than it formerly did; it is a
membranous lobe in the interior of the mouth, very conspicuous
in Orthoptera, and called the tongue, lingua, or hypopharynx ;
it reposes, in the interior of the mouth (Fig. 51, 0), on the
middle parts of the front of the labium; it is probably not
entirely lost in Coleoptera, but enters into the composition of the
STRUCTURE 97
lex middle part of the lip by amalgamation with the para-
ae. It has recently been proposed to treat this lingua as the
" ogical equivalent of the labium or of the maxillae, g giving
remains to be proved ; * the view is apparently suggested chiefly
b by the structure of the mouth of Hemimerus, a very rare and
‘most peculiar Insect that has not as yet been sufficiently studied.
: ats the maxillae and labium are largely used by taxonomists
in the systematic arrangement of the mandibulate Insects, we
sive a figure of them as seen in Coleoptera, where the parts,
though closely amalgamated, can nevertheless be distinguished.
Phie Fig. 52 should be compared with Fig. 51.
a In speaking of the segments of the body we pointed out
th they were not separate parts but constituted an uninter-
; Br ted whole, and it is well to remark here that this is also
true of the gnathites. Although the mouth parts are spoken of
? as Beerete pieces, they really form only projections from the
| body wall. Fig. 51, B, shows the intimate connexion
; “that exists between the maxillae and labium; the continuity
of the mandibles with the membrane of the bhocal cavity is
ee of very easy demonstration.
_ The head bears, besides the pieces we have considered, a pair
EG ‘antennée. These organs, though varying excessively in form,
are always present in thé adult Insect, and exist even in the
% naj ority of young Insects. They are very mobile, highly sensitive
_ organs, situate on or near the front part of the head. The
antennae arise in the embryo from the procephalic lobes, the
orphological import of which parts is one of the most difficult
“poi connected with Insect embryology.
The eyes of Insects are of two sorts, simple and compound.
dt The simple éyes, or ocelli, vary in number from one to as many
as eighteen or twenty; when thus numerous they are situated in
g oups on each side of the head. In their most perfect form, as
- found in adult aculeate Hymenoptera, in Orthoptera and Diptera,
— 0ce are usually two or three in number, and present the
pe pearance of small, perfectly transparent lenses inserted in the
integument. In their simplest form they are said to consist of
some masses of pigment in connexion with a nerve.
153
four hours a great change is found to have taken place. The
whole superficial contents of the egg are at that time arranged in
groups, having the appearance of separate rounded or oval masses,
pressed together so as to destroy much of their globular symmetry.
The egg contents are also divided into very distinct forms, a
granular matter, and a large number of transparent globules,
these latter being the fatty portion of the yolk; these are present
everywhere, though in the centre there is a space where they are
very scanty, and they also do not extend quite to the cireum-
ference. But the most remarkable change that has taken place
is the appearance in the middle of the field of an area different
from the rest in several particulars; it
oceupies about one-third of the width
and one-third of the length; it has a
whiter and more opaque appearance,
and the fat globules in it are fewer in
number and more indistinct. This
area is afterwards seen to be occupied
by the developing embryo, the outlines
of which become gradually more dis-
tinct. Fig. 83 gives an idea of the
appearance of the egg about the middle
period of the development. In warm
weather the larva emerges from this \ :
egg ten or eleven days after it has S55
been deposited. -
The period occupied by the develop- *", ae ae ears
ment of the embryo is very different in the development of the em-
the various kinds of Insects; the blowfly a en ir pat yrodby a
embryo is fully developed in less than
twenty-four hours, while in some of the Orthoptera the embryonic
stage may be prolonged through several months. According to
Woodworth the blastoderm in Vanessa antiopa is complete in
twenty-four hours after the deposition of the egg, and the
involution of the ventral plate is accomplished within three days
of deposition.
Metamorphosis.
The ontogeny, or life history of the individual, of Insects is
peculiar, inasmuch as a very large part of the development takes
154 | METAMORPHOSIS CHAP.
place only late in life and after growth has been completed. Insects
leave the egg in a certain form, and in that condition they con-
tinue—with, however, a greater or less amount of change according
to kind—till growth is completed, when, in many cases, a very great
change of form takes place. Post-embryonic development, or
change of form of this kind, is called metamorphosis. It is not a
phenomenon peculiar to Insects, but exists to a greater or less
extent in other groups of the Metazoa; while simpler post-
embryonic development occurs in nearly all, as in scarcely any
complex animals are all the organs completely formed at the time
the individual becomes possessed of a separate existence. In
many animals other than Insects the post-embryonic development
assumes most remarkable and complex forms, though there are
perhaps none in which the phenomenon is very similar to the
metamorphosis of Insects. The essential features of metamor-
phosis, as exhibited in the great class we are writing of, appear
to be the separation in time of growth and development, and the
limitation of the reproductive processes to a short period at the
end of the individual life. The peculiar phenomena of the post-
embryonic development of the white ants show that there exists
some remarkable correlation between the condition of the repro-
ductive organs and the development of the other parts of the
organisation. If we take it that the post-embryonic physio-
logical processes of any individual Insect are of three kinds,
—growth, development, and reproduction,—then we may say
that in the higher Insects these three processes are almost
completely separated, and go on consecutively, the order being,—
first, growth; second, development; third, reproduction. While,
if we complete the view by including the processes comprised in
the formation of the egg and the development therein, the series
will be—(1) oogenesis, or egg-growth ; (2) development (embry-
onic); (8) growth (post-embryonic); (4) development (post-
embryonic); (5) reproduction.
The metamorphosis of Insects is one of the most interesting
parts of entomology. It is, however, as yet very little known
from a scientific point of view, although the simpler of its
external characters have for many ages past attracted the
attention and elicited the admiration of lovers of nature. It
may seem incorrect to say that little is yet known scientifically
of a phenomenon concerning which references almost innumer-
+ sarees oo ee .
ev | METAMORPHOSIS) * 155
able are to be found in literature: nevertheless the observations
that have been made as to metamorphosis, and the analysis that -
has been commenced of the facts are at present little more than
sufficient to show us how vast and complex is the subject, and
how great are the difficulties it presents.
There are three great fields of inquiry in regard to meta-
morphosis, viz. (1) the external form at the different stages ;
(2) the internal organs and their changes; (3) the physiological
processes. Of these only the first has yet received any extensive
attention, though it is the third that precedes or underlies the
other two, and is the most important. We will say a few words
about each of these departments of the inquiry. Taking first
the external form—the instar. But before turning to this we
must point out that in limiting the inquiry to the post-embryonic
development, we are making one of those limitations that give rise
to much misconception, though they are necessary for the acquisi- *
tion of knowledge as to any complex set of phenomena. If we
assume five well-marked stages as constituting the life of an Insect
with extreme metamorphosis, viz. (1) the formation and growth of
the egg; (2) the changes in the egg culminating in its hatching
after fertilisation; (3) the period of growth; (4) the pupal
changes ; (5) the life of the perfect Insect ; and if we limit our
inquiry about development to the latter three, we are then
shutting out of view a great preliminary question, viz. whether
some Insects leave the egg in a different stage of development to
others, and we are consequently exposing ourselves to the risk
of forgetting that some of the distinctions we observe in the
subsequent metamorphosis may be consequential on differences in
the embryonic development.
Instar and Stadium.
Figs. 84 and 85 represent corresponding stages in the life
of two different Insects, Fig. 84 showing a locust (Aeridium),
and Fig. 85 a white butterfly. In each A represents the
newly-hatched individual; B, the insect just before its perfect
state ; C, the perfect or imago stage. On comparing the two sets
of figures we see that the C stages correspond pretty well as
regards the most important features (the position of the wings
being unimportant), that the A stages are moderately different,
156 ’ METAMORPHOSIS CHAP, »
while the B states are not to be recognised as equivalent condi-
tions. | :
Every Insect after leaving the egg undergoes during the
process of growth castings of the skin, each of which is called
Fic. 84. — Locust
(Acridium per-
egrinum): A,
newly hatched;
B, just -ante-
cedent to last
ecdysis ; C, per-
fect Insect.
4 - —
<= S=
J —
LSS SS =
a moult or ecdysis. Taking for our present purpose five as the
number of ecdyses undergone by both the locust and butterfly,
we may express the differences in the successions of change we
portray in Figs. 84 and 85 by saying that previous to the
Fie. 85.—Butterfly (Pieris) :
A, the newly hatched
young, or larva magnified ;
B, pupa (natural size) just
antecedent to last ecdysis ;
C, perfect Insect.
first eedysis the two, Insects are moderately dissimilar, that the
locust undergoes a moderate change before reaching the fifth
ecdysis, and undergoes another moderate change at this moult, thus
reaching its perfect condition by a slight, rather gradual series of
Vv METAMORPHOSIS fe
alterations of form. On the other hand, the butterfly under-
goes but little modification, remaining much in the condition
shown by A, Fig. 85, till the fourth, or penultimate, ecdysis,
but then suffers a complete change of form and condition, which
apparently is only inferior to another astonishing change that
takes place at the fifth or final moult. The chief, though by no
means the only, difference between the two series consists in the
fact that the butterfly has interposed between the penultimate
and the final ecdyses a completely quiescent helpless condition, in
which it is deprived of external organs of sense, locomotion, and
nutrition ; while in the locust there is no loss of these organs, and
such quiescent period as exists is confined to a short period just
at the fifth ecdysis. The changes exhibited by the butterfly are
called “ complete metamorphosis,’ while this phenomenon in the
locust is said to be “incomplete.” The Insect with complete
metamorphosis. is in its early stage called a larva, and in the
quiescent state a pupa. The adult state in both butterfly and
locust is known as imago or perfect Insect.
The most conspicuous of the differences between Insects with
complete and those with incomplete metamorphosis is, as we
have remarked, the existence in the former of a pupa. The
pupal state is by no means similar in all the Insects that
possess it. The most anomalous conditions in regard to
it occur in the Order Neuroptera. In some members of
that Order—the Caddis-flies for instance—the pupa is at first
quiescent, but becomes active before the last ecdysis; while in
another division—the May-flies—the last ecdysis is not preceded
by a formed pupa, nor is there even a distinct pupal period, but
the penultimate ecdysis is accompanied by a change of form to
the winged condition, the final ecdysis being merely a casting of
the skin after the winged state has been assumed. In the
Odonata or Dragon-flies there is no pupal stage, but the change
of form occurring at the last ecdysis is very great. In those
Insects where the interval between the last two moults is not
accompanied by the creature’s passing into a definite, quiescent
pupa, the individual is frequently called then a nymph; but the
term nymph has merely a distinctive meaning, and is not capable
of accurate definition, owing to the variety of different conditions
covered by the word. Eaton, in describing this term as it is used
for Ephemeridae, says, “Nymphs are young which lead an
158 METAMORPHOSIS CHAP.
active life, quitting the egg at a tolerably advanced stage of
morphological development, and having the mouth-parts formed
after the same main type of construction as those of the adult
insect.” + \
The intervals between the ecdyses are called stadia, the first
stadium being the period between hatching and the first ecdysis.
Unfortunately no term is in general use to express the form of
the Insect at the various stadia; entomologists say, “the form
assumed at the first moult,’ and so on. To avoid this circum-
locution it may be well to adopt a term suggested by Fischer?
and call the Insect as it appears at hatching the first instar,
what it is as it emerges from the first ecdysis the second instar,
and so on; in that case the pupa of a Lepidopteron that assumed
that condition ‘at the fifth eedysis would be the sixth instar, and
the butterfly itself would be the seventh instar.
Various terms are used to express the differences that exist in »
the metamorphoses of Insects, and as these terms refer chiefly to
the changes in the outer form, we will here mention them. As
already stated, the locust is, in our own language, said to have an
incomplete metamorphosis, the butterfly a complete one. The
term Holometabola has been proposed for Insects with complete
metamorphosis, while the appellations Ametabola, Hemimetabola,
Heterometabola, and Paurometabola have been invented for
the various forms of incomplete, or rather less complex, meta-
morphosis. Some writers use the term Ametabola for Insects |
that are supposed to exhibit no change of external form after
quitting the egg, the contrasted series of all other Insects being
then called Metabola. . Westwood and others use the word
Homomorpha for Insects in which the condition on hatching
more or less resembles that attained at the close of the develop-
ment, and Heteromorpha for those in which the form on
emergence from the egg differs much from what it ultimately
becomes.
Hypermetamorphosis.
There are certain unusual changes to which the term
hypermetamorphosis has been applied; these we can here only
briefly allude to.
1 Trans. Linn. Soc., 2nd Series, ‘‘ Zool.” 1888, iii. p. 12.
2 Orthoptera ewropaea, 1853, p. 87.
a me METAMORPHOSIS : 159
Insects that have complete metamorphoses, and are not
supplied with food by their, parents or guardians, are provided
during their larval life with special modifications of extremely
various kinds to fit them for the period of life during which they
are obtaining food and growing. Thus caterpillars possess numer-
ous adaptations to fit them for the period during which they
live on leaves, while maggots have modifications enabling them to
live amongst decomposing flesh. Some larvae are greatly modified
in this adaptive way, and when the adaptations change greatly
during the life of the larva, hypermetamorphosis is said to exist.
As an instance we may mention some beetle larvae that are born
with legs by whose aid they can clmg to a bee, and so get
Fic. 86.—Prepara-
“wh tory stages of
Sitaris humer-
alis; 9, 10, 11,
12, first, second,
third, and fourth
larval instars ;
13, pupa. (After
Lubbock and
Fabre.)
carried to its nest, where they will in future live on the stores
of food the bee provides for its own young. In order that they
may be accommodated to their totally different second circum-
stances, they change their first form, losing their legs, and _be-
coming almost bladder-like creatures, fitted for floating on the
honey without being injured by it. Such an occurrence has
been described by Fabre’ in the case of Sitaris humeralis, and
his figures have been reproduced in Sir John Lubbock’s book on
the metamorphoses of Insects,’ as well as in other works, yet they
are of so much interest that we give them again, especially as the
subject is still only in its infancy ; we at present see no sufticient
reason for the later of these larval states. Little is, we believe,
known as to the internal anatomy of the various instars in these
curious cases.
1 Ann. Sci. Nat. Zool. Ser. iv. vol. vii. 1857, pl. 17. 2 Nature Series, 1874.
160 METAMORPHOSIS CHAP.
There are certain minute Hymenoptera that deposit their
eggs inside the eggs of other Insects, where the beings hatched
from the parasitic eggs subsequently undergo their development
and growth, finding their sustenance in the yolk or embryo con-
tained in the host-egg. It is evident that such a life is very
anomalous as regards both food and the conditions. for respira-
tion, and we consequently find that these tiny egg-parasites go
through a series of changes of form of a most remarkable
character.’ It would appear that in these cases the embryonic
and post-embryonic developments are not separated in the same
way as they are in other Insects. We are not aware that any
term has yet been proposed for this very curious kind of Insect
development, which, as pointed out by Brauer, is doubtless of a
different nature from the hypermetamorphosis of Sitaris.
Changes in Internal Organs.
In relation to the post-embryonic development of the internal
organs of the body there is but little exact generalisation to be
made, the anatomical condition of these organs at the time of
emergence from the egg having been ascertained in but few
Insects. We know that in Holometabolous Insects the internal
anatomy differs profoundly in the larval and imaginal instars.
As to Insects with more imperfect metamorphosis very little
information exists, but it appears probable that in many no ex-
tensive distinctions exist between the newly-hatched and the |
adult forms, except in the condition of the reproductive organs.
Differences of minor -importance doubtless exist, but there is
almost no information as to their extent, or as to the periods at
which the changes occur; so that we do not know to what
extent they may be concentrated at the final ecdysis. In Insects
with perfect metamorphosis the structures of the internal organs
are, as we have said, in many cases totally different ‘in the larval
and imaginal periods of the life; but these changes are far from
being uniform in all Holometabola. The nervous system in
some cases undergoes a great concentration of the ganglia, in
others does not, and important distinctions exist in this respect
even within the limits of a single Order, such as the Coleoptera.
1 See Proctotrupidae subsequently,
2 Verh. Zool.-bot. Ges. Wien, xix. 1869, p. 839.
Vv METAMORPHOSIS 161
Some Insects take the same kind of food throughout their lives,
but many others change totally in this respect, and their organs
for the prehension and digestion of food undergo a corre-
sponding change. Butterflies suck food in the form of liquid
juices from flowers by means of a delicate and long proboscis,
while the young butterfly——the caterpillar—disdains sweets,
and consumes, by the assistance of powerful mandibles, a great
bulk of leaves. Other Holometabola undergo no such total change
of habits; the tiger-beetle, for instance, is as ferocious a con-
sumer of the juices of Insects in its young stage as it is in the
adult condition. Hence Brauer‘ divides Insects, as regards this
point, into three categories. The forms in which both the young
and adult take food by suction he calls Menorhyncha; those in
which both the imago and immature forms feed by mandibles he
calls Menognatha; while his Metagnatha consists of those insects
that take food by jaws when young, but by suction with tubular
mouths when mature. Besides these main divisions there are
some exceptional cases to which we need not here allude, our
present object being to indicate that in the Metagnatha the
digestive organs are of a very different nature in the young and
in the adult states of existence.
The internal organs for the continuance of the species are
known to be present in a rudimentary stage in the embryo, and
it is a rule that they do not attain their full development until
growth has been completed; to this rule there may possibly be
an exception in the case of the Aptera. But little information
of a comparative character exists as to the dorsal vessel and the
changes it undergoes during metamorphosis. There is con-
siderable difficulty in connexion with the examination of this
structure, but it appears probable that it is one of the organs
that changes the least during the process of metamorphosis.
The exact nature of the internal changes that occur during
metamorphosis is almost a modern subject. It is of course a
matter of great difficulty to observe and record changes that go
‘on in the interior of such small creatures as Insects, and when
the phenomena occur with great rapidity, as is frequently the
case in Insect metamorphosis, the difficulty is much increased,
Nevertheless the subject is of such great interest that it has been
investigated with a skill and perseverance that call for the
1 «Syst. Zool. Stud.” SB. Ak. Wien, Abth. 1, xci. 1885, p. 291.
VOL, V M
162 METAMORPHOSIS—IN TEGUMENT _ CHAP.
highest admiration. The greater part of the information ob-
tained refers to a single Insect, the blowfly; and amongst those
who have made important contributions to it we may mention
Weismann,’ Viallanes,? Ganin,® and Van Rees,* and it is at pre-
sent under investigation by Lowne. A good deal, too, is becoming
known about the processes in the case of the silkworm.
Integument and Ecdysis.
The integument consists of a cellular layer, usually called the
hypodermis, situated on a basement membrane. The hypo-
dermis, or layer of chitinogenous cells, excretes a matter which
remains attached to the body, forming the hard outer layer of the
skin. This layer consists of chitin and has no vitality, but
its presence no doubt exerts a very important influence on the
physiological processes of the Insect. The chitinous investment
varies much in thickness and in other properties; in some .
Insects it is hard, even glassy, so as to be difficult to pierce with
a pin, in others it is pliable, and in some very delicate. Chitin
is a substance very difficult to investigate; according to the
recent researches of Krawkow ° it may prove to be of somewhat
variable chemical composition.
After a time the hypodermis excretes a fresh supply of
chitin, and, possibly by the commencement of this process, the
older chitinous investment becomes separated and is shed. The
details have, however, not been ascertained, though their import-
ance has been suggested by Hatchett Jackson.6 The newly
exposed layer of integument is pallid, but afterwards becomes
coloured in a manner varying according to the species, the process
being possibly due to some secondary exudation permeating the
freshly exposed chitin, or modifying some part of its exterior.
Lowne informs us that in the imago of the blowfly the great
majority of the hypodermic cells themselves enter into the com-
position of the chitinous integument; and it is perhaps not a
matter for surprise that the cells should die on the completion of
their functional activity, and should form a part of the chitinous
1 Zeitschr. wiss. Zool. xiv. 1864, p. 187.
2 Viallanes, Ann. Sci. Nat., Series 6, “ Zool.” xiv. 1882.
% Unfortunately in the Russian language. * Zool. Jahrb. Abth. Anat. iii. 1888, p. 1.
5 Zeitschr. Biol., xxix. 1892, p. 177.
5 Trans. Linn. Soc. London, ‘‘ Zoology,” 2nd series, v. 1890, p. 174.
v METAMORPHOSIS 163
investment. Some writers say that the chitinous layer may be
shown to be covered by a delicate extima or outer coat.
The number of ecdyses varies greatly in Insects, but has been
definitely ascertained in only a few forms outside the Order
Lepidoptera. In Campodea Grassi says there is a single frag-
mentary moult, and in many Hymenoptera the skin that is cast
is extremely delicate, and the process perhaps only occurs twice
or three times previous to the pupal stage. In most Insects,
however, ecdysis is a much more important affair, and the whole
of the chitinous integument is cast off entire, even the linings of
_ the tracheae, and of the alimentary canal and its adjuncts being
parted with. Sir John Lubbock observed twenty-three moults
in a May-fly of the genus Cloéon, this being the maximum yet
recorded, though Sommer states? that in Macrotoma plumbea
moulting goes on as long as life lasts, even after the Insect has
attained its full size.
Some Insects get quit of a considerable quantity of matter by
their ecdyses, while in others the amount is comparatively slight.
It has been thought that the moulting is effected in order to
permit of increase of size of the Insect, but there are facts which
point to the conclusion that this is only a factor of secondary
importance in the matter. One of these is that many Insects
make their first ecdysis almost immediately after they leave the
egg; this is the case with the young larva of the blowfly, which,
according to Lowne, moults within two hours of its emergence
from the egg. We have already referred to the important sug-
gestion made by Eisig* that, since chitin is a nitrogenous sub-
stance, the ecdyses may be a means of getting rid of waste
nitrogenous matter; to which we have added that as chitin also
consists largely of carbon, its excretion may be of Importance
in separating carbonaceous products from the blood.
Metamorphosis of Blowfly.
The phenomena of’ metamorphosis are displayed to their
greatest extent in the transformations and physiological processes
of the Muscid Diptera, of which the common blowfly is an
_ + Trans. Linn. Soc. xxv. 1866, p. 491. * Zeitschr. wiss. Zool. xli. 1885, p. 712.
* “Fauna und Flora d. Golfes von Neapel,” Die Capitelliden, 1887, p. 781.
164 METAMORPHOSIS CHAP.
example. We will briefly consider the information that has been
obtained on this. subject. |
The development of the embryo in the egg of the blowfly is
unusually rapid, occupying only a period of twenty to twenty-
four hours. After its first moult the blowfly larva grows rapidly
during a period of about ten to fourteen days, during which it
undergoes moults, the number. of which appears not to be
definitely ascertained. After becoming full-fed the larva loses
its active state, and passes for a period into a condition of com-
parative quiescence, being spoken of in this state as a resting
larva. This quiet period occurs in most full-grown larvae, and
is remarkable for the great variation that may occur in its
duration, it being in many Insects subject to prolongation for
months, in some cases possibly even for years, though in favour-
able circumstances it may be very short. Lowne informs us that
in the blowfly this period of the life is occupied by very great
changes in the internal organs, which are undergoing very exten-
sive processes of destruction and rebuilding. After some days
the outer skin of the resting larva shrivels, and is detached from
the internal living substances, round which it hardens and forms
the sort of cocoon or capsule that is so well known. This
using of the cast larval skin as a cocoon is, however, limited to
certain of the two-winged flies, and perhaps a few other Insects,
and so must be considered an exceptional condition, The capsule
conceals from view a most remarkable state, known to the old
naturalist Réaumur as the “spheroidal condition,’ but called by
more recent writers the pronymph. . The pronymphal state
may be looked on as being to a great extent a return of the
animal to the condition of an egg, the creature becoming an
accumulation of soft creamy matter enclosed in a delicate skin,
This spheroidal condition, however, really begins in the resting
larva,and Van Rees and others think that the delicate membrane
enclosing the substance of the pronymph is really the hypodermis
of the integument of the larva. Although this seems probable,
from the resemblance this condition would in that case present
to the phenomena usual in ecdysis, it is not generally admitted,
and there is much difficulty in settling the point. Lowne is of
a contrary opinion, looking on the limiting membrane as a sub-
sequent formation; he calls it the paraderm. The process of
forming the various organs goes on in the pronymph, till the
Vv METAMORPHOSIS 165
“nymph” has completed its development, the creature having
then again taken on a definite form which apparently corre-
sponds to the pupa of Hymenoptera. Great doubt, however,
exists as to this equivalence, and indeed as to any exact corre-
spondence between the metamorphic stadia of different Insects,
a view which long since was expressed by Sir John Lubbock *
and Packard. The term nymph is used ini this case not because
there is any resemblance to the condition similarly named in
Insects with less complete metamorphosis, but because the term
pupa is applied to the outer case together with the contained
nymph. The transformation of the nymph into the perfect blow-
fly occupies a period very variable according to the temperature.
Histolysis.—The processes by which the internal organs of
the maggot are converted into those of the fly are of two kinds,
—histolysis or breaking down, histogenesis or building up, of
tissue. The intermediary agents in histolysis are phagocytes,
cells similar to the leucocytes or white corpuscles of the blood:
the intermediary agents in histogenesis are portions of tissue
existing in the larval state incorporated with the different organs,
or preserving a connexion therewith even when they are to a
great extent separated therefrom. In this latter case they are
called imaginal discs, though Professor Miall prefers to term them
imaginal folds.” The two processes of histolysis and histogenesis,
though to some extent mutually dependent (for the material to be
built up has to be largely obtained by previous destruction), do not
go on pari passu, though they are to a great extent contemporaneous.
In the resting larva histolysis is predominant, while in the nymph
histogenesis is more extensive. Microscopic observation shows
that the phenomena connected with the histolysis of the mus-
cular tissue are scarcely distinguishable from those of an inflam-
matory process, and Viallanes* dilates on this fact in an instruc-
tive manner. The phagocytes attach themselves to, or enter, the
tissues which are to be disintegrated, and becoming distended,
assume a granular appearance. By this pseudo-inflammatory pro-
cess the larval structures are broken down into a creamy substance ;
the buds, or germs, from which the new organs are to be devel-
oped being exempt from the destruction. These buds, of which
about sixty or upwards have already been detected, undergo
? Trans. Linn. Soc. xxiv. 1863, p. 65. 2 Trans. Linn. Soc. “Zool.” v. 1892, p. 267.
. 3 Ann. Sci. Nat., Series 6, ‘‘ Zool.” xiv. 1882, p. 150.
166 METAMORPHOSIS CHAP,
growth as they are liberated, and so the new creature is formed,
the process of growth in certain parts going on while destrue-
tion is being accomplished in others. Considerable discrepancy
prevails as to the extent to which the disintegration of some
of the tissues is carried.
According to Kowalevsky* it would appear that after the
phagocytes have become loaded with granules they serve as
nutriment for the growing tissues, and he thinks they become
blood-cells' in the imago. The process of histolysis has been
hy af
V Ge jj :
{i}
fi
Wy
Ng =
4 \
?
if EEE
L) {
j ia
\
Fie, 87.—Imaginal discs of Muscidae in process of development: A, Brain and
ventral ganglion of a larva 7 mm. long of M. vomitoria; v, ventral ganglion ;
c, cephalic ganglion; h, head rudiment; ve, portion of: ventral chain; pd,
prothoracic rudiment; vc3, third nerve; md, mesothoracic rudiment: B, meso-
thoracic rudiment, more advanced, iu a pupa just formed of Sarcophaga carnaria,
showing the base of the sternum and folds of the forming leg, the central part (/)
representing the foot: C, the rudimentary leg of the same more advanced ; f, femur;
t, tibia ; /,, 7; tarsal joints: D, two discs from a larva 20 mm. long of Sarcophaga,
attached to tracheae ; msw, mesonotal and wing-rudiment ; m¢, metathoracie rudi-
ment; E, 7, mesothoracic rudiment of a 7 mm. -long larva attached to a tracheal
twig. (After Weismann and Graber. )
chiefly studied in the blowfly, and not much is known of it in
other Insects, yet it occurs to a considerable extent, according to
Bugnion” and others, in the metamorphosis of Lepidoptera.
Indeed it would almost seem that the processes of histolysis
and histogenesis may be looked on as exaggerated forms of the
phenomena of the ordinary life of tissues, due to greater rapidity
and discontinuity of tissue nutrition. |
1 Zool. Anz. viii. 1885, p. 125. 2 Mitt. Schweiz. ent. Ges. viii. 1893, p. 403.
’ ,
‘
Py
«
ee
at a Eee
~ METAMORPHOSIS 167
_ Imaginal Discs.—The imaginal discs are, portions of the
larval hypoderm, detached from continuity with the main body
of the integument, but connected therewith by strings or pedicels
which may be looked on as portions of the basement membrane.
Whether these discs, or histoblasts as they are called by Kiinckel
d’Herculais,’ are distinguished by any important character from
other buds or portions of regenerative tissue that, according to
Kowalevsky,? Korschelt and Heider,? and others, exist in other
parts of the body, does not appear to be at present ascertained.
We give some figures, taken from Weismann and Graber, of
- the imaginal rudiments existing in the
larvae of Muscidae. Although by no means
good, they are the best for our purpose
we can offer to the reader. Other figures
will be found in Lowne’s work on _ the
blowfly now in course of publication.
Weismann’s paper * is now thirty years old,
and, when it was written, he was not aware
of the intimate connexion the rudiments
have with the integument; this has, how-
ever, now been demonstrated by several
observers. Pratt states’ that the formation
of the imaginal discs in Melophagus ovinus
takes place in the later stages of the em-
bryonic development, and after the manner
formerly suggested by Balfour, viz. invagin-
ation of the ectoderm.
Both the regenerative buds and the
‘rudimentary sexual glands are known to be
derived directly from the embryo; neither
of them undergoes any histolysis, so that Fie. 88.— Median longi-
we have in them embryonic structures ‘inal section through
: Seman : fo , larva of blowfly during
which exist in a quiescent condition during _ the process of _histo-
th ‘od i Roy the. } ey . lysis. (After Graber.)
e period in which the larva is growing — fxpjanation in text.
with great rapidity, and which when the
larva has attained its full growth and is disintegrating, then
1 Recherches Org. des Volucelles, 1875, p. 143.
2 Zeitschr. wiss. Zool. xlv. 1887, p. 587.
3 Lehrbuch Entwicklungsgeschichte, Spec. Theil. 1890, p. 875.
* Zeitschr. wiss. Zool. xiv. 1864, p. 187. ° Arch. f. Naturges. lix. 1893, 1, p. 168.
168 METAMORPHOSIS CHAP, Om
appropriate the products of the disintegration so as to’ produce
the perfect fly.
Our Fig. 88, taken from Graber, represents a longitudinal
median section of a full-grown larva of Musca, in which the
processes of metamorphosis are taking place. The position of
some of the more important imaginal rudiments is shown by it:
b', 6, 8, rudiments of the three pairs of legs of the imago; an, of
antennae ; between an and w, rudiment of eye; w, of wings; h, of
halteres ; f, fat-body ; d, middle of alimentary canal; n, ventral
chain; st, stigma; 6, 7, sixth and seventh body segments.
Physiology of Metamorphosis.
Many years ago, Harvey perceived the probable existence of
a physiological continuity between the earlier and later stages
of the Insect’s life. Modern investigation has shown that in the
blowfly a remarkable analogy exists between the conditions of
the pupa and the egg. The outer shell of the pupa corresponds
to the chorion or egg-shell, and the delicate outer membrane of
the pronymph to the oolemn or lining membrane of the egg; the
creamy matter corresponds with the yolk, and the regenerative
buds are analogous to the formative portions of the developing
egg. The process of histolysis as carried out by the phagocytes
of the later life appears also to find a parallel in the vitellophags
of the embryonic life.’ It appears probable that the physio-
logical processes of the post-embryonic metamorphosis may be
essentially a repetition—or an interrupted continuation—of
those of the embryonic period.
The inquiry as to what are the determining causes of the
metamorphic changes of the blowfly and other Insects has as
yet but little advanced. Why does the larva grow up to a
certain period with great rapidity, then cease its appropriating —
power and break up the parts that have been so rapidly and
recently formed? And why do the imaginal buds remain
quiescent till the other tissues are being disintegrated, and
then, instead of sharing the general condition of disintegra-
tion, commence a career of development? To these questions no
satisfactory answer has yet been given, though the remarkable
- studies, already referred to, of Bataillon on the later larval life
1 Wheeler, in J. Morphol. viii. 1893, p. 81.
Ee ee ee ee A ae reer Mn Ee aa i ae OR ae a
METAMORPHOSIS 169
of the silkworm suggest the direction in which knowledge may
be found, for they show that the physiological conditions of the
later larval life are different from those of the earlier life, possibly
as the direct result of the mere aggregation of matter, and the
consequent different relations of the parts of the organism to
atmospheric and aqueous conditions.
If we wish to understand metamorphosis, we must supplement
the old opinion that ecdysis is merely an occurrence to facilitate
expansion, by the more modern conception that it is also an
important physiological process. That shedding the skin is done
solely to permit of enlargement of size is a view rendered unten-
able by many considerations. The integument can increase and
stretch to an enormous extent without the aid of moulting; wit-
ness the queen-termite, and the honey-bearers of the Myrmeco-
cystus ants. Many moults are made when increase of size does
not demand them, and the shedding of the skin at the time of.
pupation is accompanied by a decrease in size. And if moulting
be merely connected with increase of size, it is impossible to see
why Cloéon should require two dozen moults, while Campodea
can do with one, or why a collembolon should go on moulting
during the period of life subsequent to the cessation of growth.
The-attention of entomologists has been chiefly directed to
the ecdyses connected with the disclosure of the pupal and
imaginal instars. Various important transformations may, how-
ever, occur previous to this, and when they do so it is always
in connexion with ecdyses. Caterpillars frequently assume a
different appearance and change their habits or character at a
particular ecdysis; and in Orthoptera each ecdysis is accom-
panied by a change of form of the thoracic segments; this
change is very considerable at one of the intermediate ecdyses.
The assumption of the pupa state is the concomitant of an
ecdysis, and so also is the appearance of the imago; but the
commencement of each of these two stages precedes the ecdysis,
which is merely the outward mark of the physiological processes.
The ecdysis by which the pupa is revealed occurs after the
completion of growth and when great changes in the internal
organs have occurred and are still taking place; the ecdysis by
which the imago appears comes after development has been quite
or nearly completed.
Although the existence of a pupa is to the eye the, most
170 _-—— *_s METAMORPHOSIS CHAP. v.
striking of the differences between Insects with perfect and those
with imperfect metamorphosis, yet there is reason for supposing
that the pupa and the pupal period are really of less importance
than they at first sight appear to be. In Fig. 85 we showed
how great is the difference in appearance between the pupa and
the imago. The condition that precedes the appearance of the
pupa is, however, really the period of the most important change.
In Fig. 89 we represent the larva and pupa of a bee; it will be
seen that the difference between the two forms is very great,
while the further change that will be required to complete
the perfect Insect is but slight. When the last skin of the
Fria. 89.—Larva and pupa of a bee, Xylocopa violacea: A, larva; B, pupa, ventral ©
aspect ; C, pupa, dorsal aspect. (After Lucas.) °
larva of a bee or of a beetle is thrown off, it is, in fact, the
imago that is revealed; the form thus displayed, though colour-
less and soft, is that of the perfect Insect; what remains to be
done is a little shrinking of some parts and expansion of others,
the development of the colour, the hardening of certain parts.
The colour appears quite gradually and in a regular course,
the eyes being usually the first parts to darken. After the
coloration is more or less perfected—according to the species
—a delicate pellicle is shed or rubbed off, and the bee or beetle
assumes its final form, though usually it does not become active
till after a farther period of repose.
CHAPTER VI.
CLASSIFICATION——THE NINE ORDERS OF INSECTS—-THEIR CHARACTERS
—PACKARD’S ARRANGEMENT——BRAUER’S CLASSIFICATION—
CLASSIFICATIONS BASED ON METAMORPHOSIS——SUPER-ORDERS
—THE SUBDIVISIONS OF ORDERS.
Classification.
WE have already alluded to the fact that Insects are the most
numerous in species and individuals of all land animals: it is
estimated that about 250,000 species have been already described
and have had scientific names given to them, and it is considered
that this is probably only about one-tenth of those that really
exist. The classification in a comprehensible manner of such an
enormous number of forms is, it will be readily understood, a
matter of great difficulty. Several methods or schemes have
‘since the time of Linnaeus been devised for the purpose, but we
shall not trouble the reader to consider them, because most of
them have fallen into disuse and have only a historical interest.
Even at present there exists, however, considerable diversity of
opinion on the question of classification, due in part to the fact
that some naturalists take the structure of the perfect or adult
_ Insect as the basis of their arrangement, while others prefer to
treat the steps or processes by which the structure is attained, as
being of primary importance. To consider the relative values of
these two methods would be beyond our scope, but as in practice
a knowledge of the structures themselves must precede an inquiry
as to the phases of development by which the structures are reached;
and as this latter kind of knowledge has been obtained in the
case of a comparatively small portion of the known forms,—the
embryology and metamorphosis having been investigated in but
172 INSECTS CHAP,
few Insects,—it is clear that a classification on the basis of
structure is the only one that can be at present of practical value.
We shall therefore for the purposes of this work make use of an
old and simple system, taking as of primary importance the nature
of the organs of flight, and of the appendages for the introduction
of food to the body by the perfect Insect. We do not attempt to
disguise the fact that this method is open to most serious
objections, but we believe that it is nevertheless at present the
most simple and useful one, and is likely to remain such, at any
rate as long as knowledge of development is in process of
attainment.
Orders.
The great groups of Insects are called Orders, and of
these. we recognise nine, viz. (1) Aptera, (2) Orthoptera, (3)
Neuroptera, (4) Hymenoptera, (5) Coleoptera, (6) Lepidoptera,
(7) Diptera, (8) Thysanoptera, (9) Hemiptera. These names are
framed to represent the nature of the wings; and there is some
advantage in having the Orders named in a uniform and descriptive
manner. The system we adopt differs but little from that
proposed by Linnaeus." The great Swedish naturalist did not,
however, recognise the Orders Orthoptera and Thysanoptera ; and
his order Aptera was very different from ours.
being asked to recall the fact that by a mandibulate mouth we
understand one in which the mandibles, or the maxille, or
both, are fitted for biting, crushing, or grasping food; while the
term suctorial implies that some of the mouth parts are of a
tubular form or are protrusible as a proboscis, which assists, or
protects, a more minute and delicate sucking apparatus :—
1. Aptera (a without, rrepdv a wing). Wingless? Insects ; mouth mandibulate
or very imperfectly suctorial. Metamorphosis very little.
2. Orthoptera (opOds straight, rrepov a wing). Four wings are present, the
front pair being coriaceous (leather-like), usually smaller than the
other pair, which are of more delicate texture, and contract in repose
after the manner of a fan. Mouth mandibulate. Metamorphosis
slight. ;
3.“ Neuroptera (vetpov nerve, rrepdv a wing). Four wings of membranous
1 Syst. Nat. Ed. 12, ref. i. pars ii. p. 536 (by error, 356).
2 It must not be supposed that all wingless Insects fall within the limits of this .
Order.
ail
VI _ THE ORDERS OF INSECTS 173
consistency, frequently with much network ; the front pair not much,
if at all, harder than the other pair, the latter with but little or no
fanlike action in closing,- Mouth mandibulate. Metamorphosis
variable, but rarely slight.
4, Hymenoptera (ipjv membrane, rrepov a wing), Four wings of membranous
consistency ; the front pair larger than the hind, which are always
small and do not fold up in repose. Mouth mandibulate, sometimes
provided also with. a tubular proboscis, Metamorphosis very great.
5. Coleoptera (koXeds sheath, rrepdv a wing). Four wings; the upper pair
shell-like in consistency, and forming cases which meet: together
over the back in an accurate line of union, so as to entirely lose a
° winglike appearance, and to conceal the delicate membranous hind
pair. Mouth mandibulate. Metamorphosis great.
6. Lepidoptera (Aeris scale, trepdv a wing). Four large wings covered with
scales. Mouth suctorial. Metamorphosis great.
7. Diptera (Sis double, rrepdv a wing), Two membranous wings. Mouth
suctorial, but varying greatly. Metamorphosis very great.
8. Thysanoptera (Ovoavos fringe, rrepdv a wing). Four very narrow fringed
wings. Mouth imperfectly suctorial. Metamorphosis slight.
9. Hemiptera (pe half, rrepdv a wing). Four wings; the front pair either
leather-like with more membranous apex, or entirely parchment-like
or membranous. Mouth perfectly suctorial, Metamorphosis usually
slight.
We must again ask the reader to bear in mind that numerous
exceptions exist to these characters in most of the great Orders ;
for instance, wingless forms are not by any means rare in several
of the Orders.
Before remarking further on this system we will briefly
sketch two other arrangements of the Orders of Insects, for which
we are indebted to Packard and Brauer.
Packard’s Classification.
Packard has devoted much attention to the subject, and has
published two or three successive schemes, of which the following
is the most recent:’ the definitions are those of the author
himself, but the information in brackets is given to institute a
concordance with the system we adopt :— -
1. Thysanura. Wingless ; often with a spring (equivalent to our Apftera).
2. Dermaptera. Front wings minute, elytra-like (= Forficulidae, a part of
our Orthoptera).
3. Orthoptera. Wings net-veined ; fore wings narrow, hind wings folded
(=our Orthoptera after subtraction of Dermaptera).
1 American Naturalist, xx. 1886, p. 808.
?
174 INSECTS CHAP, — |
+
4. Platyptera. Four net-veined wings; mouth parts adapted for biting
(= Termitidae and Mallophaga, parts of our Neuroptera).
5. Odonata. Wings net-veined, equal (= Odonata, a division of our
Neuroptera).
6. Plectoptera. Wings net-veined, unequal (= Hphemeridae, a part of our
Neuroptera).
7. Thysanoptera. Mouth beaklike but with palpi (= our Thysanoptera).
8. Hemiptera. Mouth parts forming a beak for sucking. No palpi (=our™
Hemiptera). ;
The above eight Orders form the group AMETABOLA, while the
following eight constitute the METABOLA :— .
9. Neuroptera. Wings net-veined ; metamorphosis complete (=a small part
of our Neuroptera).
10. Mecaptera. Wings long and narrow (fora small part of our Neuroptera;
the Panorpatae of Brauer).
11. Trichoptera. Wings not net-veined (=our division of Newroptera with
the same name). .
12. Coleoptera. Fore wings sheathing the hinder ones (= our Coleoptera).
13. Stphonaptera. Wingless, parasitic. Flea (=a division of Diptera).
14, Diptera. One pair of wings (=our Dvzptera after subtraction of
Siphonaptera).
15. Lepidoptera, Four wings (and body) scaled (= our Lepidoptera).
16, Hymenoptera. Four clear wings; hinder pair small; a tongue (=our
Hymenoptera). . |
Se te on 6 ee
Although this system of the Orders of Insects has some
valuable features it 1s open to very serious objections, to which we
can only briefly allude. The Order Hemiptera with its extensive —
divisions, Heteroptera, Homoptera, Coccidae, and Anoplura exhibit- _
ing great differences in structure and considerable divergence in
metamorphosis, is treated as only equivalent to the little group
Panorpatae (scorpion-flies); these latter being considered a dis-
tinct order, although they are not very different in structure or
metamorphosis from the Orders he calls Neuroptera and Trichop-
tera. The arrangement appears to be specially designed with
the view of making the Orders adopted in it fall into the two
groups Ametabola ‘anid Metabola. The propriety of such a
course is more than doubtful since very few of the Ametabola
are really without metamorphosis, in the wide sense of that term,
while the Metabola include Insects with various kinds of meta- .
morphosis. Indeed if we substitute for the term Ametabola the
more correct expression, “ Insects with little metamorphosis,” and
for Metabola the definition, “Insects with more metamorphosis
but of various kinds,” we then recognise that the arrangement
vI CLASSIFICATION - 175
is, like all others, a quite artificial one, while it is of little
value, owing to the development of so few Insects being hitherto
fully ascertained.
Brauer’s Classification.
Professor Brauer has recently proposed * to adopt 17 Orders
or chief groups of Insects, arranging them as follows :—
I, APTERYGOGENEA (with one order).
1. Synaptera (= Aptera of our system).
IL PreryeocEenra (=all the other Insects of our arrangement).
2. Dermaptera (= Orthoptera, Fam. Forficulidae in our arrangement).
3. Ephemeridae (=a division of Neuroptera in our arrangenient).
4, Odonata (=a division of Neuroptera in our arrangement).
5. Plecoptera (= Neuroptera, Fam. Perlidae in our arrangement).
6. Orthoptera (= our Orthoptera — Forficulidae and + Embiidae).
7. Corrodentia (=the families Termitidae, Psocidae, and Mallophaga, of
our Neuroptera).
8. Thysanoptera (as with us).
9. Rhynchota (= Hemiptera with us).
10. Neuroptera (= the families Hemeroliide and Sialide of our Neuroptera),
11. Panorpatae (=the family Panorpidae of our Neuroptera).
12. Trichoptera (=the division Trichoptera of Neuroptera).
13. Lepidoptera (=as with us).
14. Diptera (= our Diptera — Aphaniptera).
15. Siphonaptera (= Aphaniptera, a division of Diptera with us).
16. Coleoptera (= Coleoptera).
17. Hymenoptera (as with us).
The chief characters on which Brauer bases his system are:
(1) The existence or absence of wings. (2) The condition of the
mouth, and whether it undergoes radical changes.in the ontogeny,
arriving thus at the categories Menognatha, Metagnatha, and
Menorhyncha, as we have mentioned on p. 161. (3) The meta-
morphosis; the grouping adopted being Ametabola, Hemimetabola,
Metabola. (4) The number of the Malpighian tubules ;
Oligonephria, Polynephria. (5) The nature of the wings, the
relative proportions of the thoracic segments, and some other
characters. |
Brauer’s treatise is accompanied by a valuable and in many
respects very sagacious consideration of the generalised char-
acters of the Insecta; as a classification based partly on general-
isations and partly on structures, it is, so far as the present
1 “Syst. Zool. Studien.” §.B. Ak. Wien, xci. 1885, Abth, I. p. 374.
176 CLASSIFICATION CHAP.
condition of our knowledge goes, a good one. But it is of
little value as a practical guide, and as a basis for theoretical
speculation it cannot be treated as of importance, because the
generalisations it makes use of are premature, owing to the small
proportion of the forms that have been examined. And even now
the groups adopted are known to be subject to many exceptions.
Thus it begins by a division of Insecta into. winged and
wingless; but the winged division is made to comprehend an
enormous number of wingless Insects, whole subdivisions of
Orders such as the Mallophaga being placed in the winged series,
although all are without wings. This first division is indeed
entirely theoretical; and if a classification on generalisations
were adopted, it would be more natural to begin with the old
division into Homomorpha and Heteromorpha, and treat the
Order Aptera as the first division of the Homomorpha, while the
Heteromorpha would commence with the Ephemeridae and Odonata,
in which, though the individual in the early part of the ontogeny
is very different from the perfect Insect, there is no marked
division of the later larval and the pupal stages. Brauer’s system
is also defective inasmuch as it takes no account of the embryo-
logical or oogenetic processes, though these are of equal import-
ance with the later phases of the Ontogeny, Even as regards the
division into Orders, it is far from being free from reproach ; for
instance, the separation of the Dermaptera from the Orthoptera,
while Rhynchota remains intact, although including a more
extensive series of heterogeneous forms; the division of the-
Neuroptera into widely separated groups, each of which is treated
as equivalent to the great Orders, such as Coleoptera (in which
Strepsiptera are included), Hymenoptera, and Diptera, is not
reasonable, The association of Mallophaga and Termitidae, while
Dermaptera are separated from Orthoptera, is also undeniably
arbitrary, and other similar disparities are to be seen on
scrutinising the details of the system. 7
On comparing the three arrangements we have outlined, it
will be seen that the chief discrepancies they present come
under two heads: (1) The treatment of the Neuroptera, opinions
differing as to whether these Insects shall be grouped as a single
Order, or shall be divided into numerous Orders; and as to what,
if this latter course be adopted, the divisions shall be. (2) The
treatment of the parasitic groups Mallophaga, Aphaniptera, ete.
VI CLASSIFICATION 177
It must be admitted that whichever of the three systems we
have sketched be adopted, the result is, as regards both these
points, open to criticism. The Order Neuroptera, if we take it
in the broad sense, differs from the other Orders in the greater
variety of metamorphosis exhibited by its members; while if, on
the contrary, it be dismembered, we get a number of groups
of very unequal extent and not distinguished from one another
by the same decisive and important characters as are the other
Orders of which they are considered equivalent. The discrepancy
exists in nature, and can scarcely be evaded by any system.
eernnenggea Maree har we 29 6
ae ;
VII THYSANURA 185
- Blattidae; Grassi states, however, that not only are they eyes,
but that they are of almost unique structure, being, in fact,
intermediate between simple and compound eyes.
The mode of development of the compound eyes of Machilis
is of considerable interest, but unfortunately very little is known
about it, even the period at which the eyes appear being uncertain.
Judging from analogy with the Orthoptera, we should suppose
them to be present when the Insect leaves the egg, and Oudemans
apparently considers this to be the case, but Bolivar states’ that
Fie. 92.—Head of Machilis mari-
tima (after Oudemans): A, base
of antenna; C, clypeus; /, ver-
tex; P, fold; O, eye; 0, 0’, sup-
posed simple eye; J/, mandible ;
m, maxilla; L, upper lip; 7, lower Fie. 93.—Lepisma cineta. (After
lip; 7, portion of maxillary palp ; Oudemans.) x4. (The line indi-
t, of labial palp. x 20. cates the natural length, )
in the early stages of Machilis the eyes are only simple eyes;
these being replaced by compound eyes in the later life. The
writer has observed very young individuals of, Machilis polypoda,
and found the eyes to be evidently compound.
The remaining family of Thysanura, the Lepismidae, is in
certain respects the most highly developed of the Order. The
covering of scales found on the body is very remarkable in some
of the species, especially in the genus Lepisma (Fig. 93, J.
cincta); the thoracic segments are different from one another
1 Ann. Soc. ent. France, 1892, p. 34.
186 APTERA © s OHAP.
and from those of the abdomen, and the tracheal system is more
highly developed than it is in the Machilidae. Several genera
are known, but only two members of the family have yet been
detected in Britain. One of them (Lepisma saccharina), occurs
only in houses, and is sometimes called the silver fish; it is,
when full grown, less than half an inch long, and is covered with
scales that give it a feebly metallic lustre. Like the other
Thysanura, its movements are very perfect. It is said that it is
occasionally injurious by nibbling paper, but the writer’s observa-
tions lead him to doubt this; its usual food is doubtless farin-
aceous or saccharine matter. Zhermobia furnorum, our other
British Lepismid, has only recently been discovered ; it is found
in bakehouses at Cambridge and elsewhere. The bakers call
these Insects fire-brats, apparently considering them to be fond
of heat. |
Much valuable information as to the anatomy of Thysanura
has been obtained by Grassi and Oudemans, and is of great
interest. Taking four genera, viz. Campodea, Japyx, Machilis,
and Lepisma, to represent the four families constituting the
sub-order, we will briefly enumerate some of the more remarkable
of the characters of their internal anatomy. Campodea has a
very inferior development of the tracheal system; there are three
pairs of spiracles, which are situate on the thoracic region;
the tracheae connected with each spiracle remain distinct,
not uniting with those coming from another spiracle; there are
thus six separate small tracheal systems, three on each side
of the body. Japyx solifugus has eleven pairs of spiracles, of
which four are thoracic; the tracheae are united into one system
on each side by means of lateral tubes; thus there are two
extensive tracheal systems situate one on each side of the body,
there being a single transverse tube, placed near the posterior
extremity, uniting the two lateral systems. In Machalis there
are nine pairs of stigmata, two of them thoracic, seven abdominal ;
the tracheae from each spiracle remain unconnected, so that there
are eighteen separate tracheal systems, some of which are con-
siderably more developed than others. The Lepismidae have
ten pairs of stigmata, and the tracheae connected with them are
completely united into one system by longitudinal and transverse
tubes. Besides these differences there are others, of considerable
importance, in the position of the stigmata.
THYSANURA 187
_ All the Thysanura possess salivary glands. In Campodea
there are about sixteen extremely short Malpighian tubules, or
1 ) Sena glands representing these organs; Japyxz is destitute
' these structures; Machilis maritima has twenty elongate
tubu es; 1n Pee also they are long, and apparently vary in
mber from four to eight in different species. The propor-
tions of the three divisions of the alimentary canal differ’
extremely ; there is a very large proventriculus in Lepisma, but
ot ot in the other families; coecal diverticula are present on the
me rior part of the true Honiadh in Machilis and in Lepisma,
- j are wanting in Campodea and in Japyz.
The dorsal vessel seems not to present any great differences
n the sub-order. Grassi says there are no alary muscles present,
ie Oudemans describes them as existing in Machilis, but as
sing excessively delicate.
i The ventral chain of nerve-ganglia consists in Campodea of
: ne cephalic ganglion, one sub-oesophageal (which clearly
belongs to the ventral series of ganglia), three thoracic, and
seven abdominal. In the other families there are eight instead
of seven abdominal ganglia.
ag structure of the internal sexual organs is very remark-
able in the Thysanura. In Campodea there is one extremely
a ze, Simple tube on each side of the body. In Japys there are
‘seven small tubes on each side, placed one in each of the suc-
cessive abdominal segments, and opening into a common duct.
; _ Machilis there are also seven tubes opening into a common
luct, but the arrangement is no longer a distinctly segmental one.
in Lepisma there are five egg-tubes on each side, the arrange-
aent being segmental in the young state but not in the adult
condition. In Campodea nutrient cells alternate with the eggs
n the tubes, but this is not the case in the other families,
Fi ig. 94 shows the ovaries in three of the Thysanura ; in the draw-
ing representing this part in Machilis (C), one of the two ovaries
is cut away for the sake of clearness.
The male organs in Campodea are very similar in size and
oo to the ovaries, there being a single large tube
n° * 8aC and a short vas deferens on each side of the body. In
yx there is a sac on each side, but it is rendered double by
, coeeum at its base, and there are long and tortuous vasa
entia. In ZLepisma there are three pairs of coeca on each
188 THYSANURA ALS CHAP.
side, segmentally placed and opening into a common duct.
In Machilis there are three retort-shaped sacs on each side open-
ing near one another into a common duct, the vasa deferentia
are elongate, and are very curiously formed, being each double
for a considerable length, and the separated portions connected
at intervals by five transverse commissural ducts.
One of the characteristic features of Insect structure is the —
restriction of articulated legs to the thoracic region. In the
Thysanura there exist ap-
pendages occupying a posi-
tion on the hind body some-
what similar to that of the
legs on the thorax. These
appendages are quite small
bodies, and are placed at the
C hind margins of the ventral
| plates of the abdomen, one
near each side; they are con-
nected by a simple joint to
the sternite and are provided
with muscles. They are
found in Campodea on seg-
ments 2 to 7; in Lepisma
| on 8 and 9, in the allied
Fic. 94.—Ovaries of Thysanura: A, of Cam- Nicoletia on 2 to 9; in Japyx
podea } B, of Jupyx ; C,of Machilis. (After on. dete 7, being, how- —
Grassi and Oudemans. ) ;
ever, more rudimentary than
they are in Campodea. In Machilis they attain perhaps their
greatest development and exist on segments 2 to 9; more-
over, in this genus such appendages occur also on the coxae of
the second and third pairs of thoracic legs. Oudemans thinks
they help to support the abdomen, and that they also assist
in leaping; Grassi considers that they are supporting agents to
some extent, but that they are essentially tactile organs. He
calls them false legs “ Pseudozampe.”
Still more ‘omarion and obscure in function are the vesicles
found near the appendages; we figure a pair after Oudemans,
showing them in the exserted state. In the retracted state the
outer portion of the vesicles is withdrawn into the basal part P
(Fig. 95), so that the vesicles are then only just visible, being
vir | APTERA 189
concealed by the ventral plate. The abdominal appendage is not
retractile. In Machilis there are twenty-two of these vesicles,
arranged either two or four on one ventral plate of the hind
body. They are also present in Campodea, where there are six
pairs. They are usually said to be absent in Japyx and in
Lepisma, but Haase shows' that Japya possess a pair placed
behind the second ventral plate of the abdomen. The vesicles
appear to be exserted by the entrance of blood into them, and to
be retracted by muscular agency. Much difference of opinion
prevails as to their function; it appears probable that they may
be respiratory, as suggested by Oudemans.
The scales found on the bodies of the Ectotrophous Thysanura
may be looked on as modified hairs, and are essentially similar to
those of the Lepidoptera, and they drop off
as readily as do those of the Lepidoptera.
Stummer-Trauntels, who has recently
published * the results of his researches on
the mouth-organs of Thysanura and Col-
lembola, confirms the division of the
Thysanura into Entotrophi and Ectotrophi,
and considers that the Collembola agree
with the former group. The German
author therefore proposes to divide our
Aptera, not into Thysanura and Collembola, gr Aion aaa aa
but into Ectognathi and Entognathi, the cles of Machilis. A,
former group consisting of Machilidae and sion te ig ae
Lepismidae, the latter of Campodeidae, tion; &, muscles. x70.
Japygidae and the various families of
Collembola. We think it far more natural, however, to retain
the older division into Thysanura and Collembola.
Collembola.
The sub-order Collembola, which we have defined on p. 182,
consists of small Insects, many of which possess the capacity of
leaping, or springing suddenly, and when disturbed or alarmed
naturally make use of this means of escaping. Their leaps, how-
ever, appear to be made quite at random, and very frequently do
* Morph. Jahrb. xv. 1889, p. 363. 2 SB. Ak. Wien, c. 1891, Abth. I. p. 216.
190 COLLEMBOLA . CHAP.
not have the result of taking the creature into concealment, and
in such circumstances they may be rapidly and frequently
repeated until the Insect feels itself, as we may suppose, in a
position of safety. Three families may be very readily. dis-
tinguished, viz. (1) Lipuridae, in which no leaping apparatus is
present; (2) Poduridae, a leaping apparatus exists near the ex- —
tremity of the abdomen; the body is subcylindric and evidently
segmented; (3) Smynthuridae, a leaping apparatus exists: the ~
body is sub-globular with comparatively large head and abdomen,
the intervening thoracic region being small; the segmentation of
the body is obscure.
The study of the Collembola is much less advanced than that
of the Thysanura, comparatively little having been added to our
knowledge of the group since Lubbock’s monograph of the
British forms was published by the Ray Society in 1873. Why
the Collembola should be neglected when the Thysanura attract
so much attention is as inexplicable as many other fashions are.
The family Lipuridae consists of a few very small and obscure
Insects of soft consistence. They move slowly, and, owing to the
absence of any leaping power, attract atten-
tion less readily than the other Collembola
do. ‘Two genera are generally recognised,
and they should probably form separate
families ; indeed, in Lubbock’s arrangement
they do so. In one of the genera (Anoura)
the mouth is very imperfect, no mandibles
or maxillae having been detected, while in
the other genus (Lipura) these organs exist.
In the members of the family Poduridae,
including the Degeeriidae of Lubbock, a
saltatory apparatus is present in the form
of avpendages attached to the fifth abdomi-
nal segment (Degeeriides), or to the fourth
Fic, 96.—Lipura bur- (Podurides). These appendages are during
eerie i. (After Lub- Jife flexed beneath the body, but in dead
specimens usually project backwards, having
the appearance of a bifid tail. Poduridae are of elongate form,
somewhat like small caterpillars, and are frequently prettily
marked with variegate colours. Fig. 97 represents an arctic form
closely allied to our native genus Jsotoma.
vit SPRING-TAILS IgI
The peculiar shape of the members of the Smynthuridae is
sufficient for their identification. They possess a very convex
abdomen, and very near to it
a large head, the intervening
chink being occupied by the
small thorax. The segmenta-
tion of the body is not easily
distinguished. Nicolet states
that the thorax consists of
three segments and the abdo-
men of the same number, and
that when the Insect emerges Fic. 97.—Corynothrix borealis: a, ventral
Peer the ege Me, divisions tube ; 6, the spring. (After Tullberg.)
can be perceived. In after life the posterior part of the thorax
becomes amalgamated with the abdomen, so that it is difficult to
trace the divisions, but there appears to be no information as to
- the manner in which this change occurs.
Some of these minute Insects frequent
trees and bushes, and their leaping powers
are very perfect, so that it is difficult to
capture them. The family includes both
the Smynthuridae and the Papiriidae of
Lubbock.
The two most characteristic organs of
the Collembola are the spring and the
ventral tube. The first of these is an
elongate structure attached to the under-
side of the abdomen near its extremity,
either on the penultimate or ante-penulti-
mate segment. It consists of a_ basal
part, and of two appendages attached
thereto. It is carried under the Insect
bent forwards, and is retained in this
position by means of a catch which pro-
jects from the under surface of the third
Fig. 98.—Smynthurus varie. Segment of the body, descending between
Oe, ation Tabs a) the two branches of the spring, and pass-
ing under the extremity of its basal seg-
ment. It is considered that the spring is elastic, is flexed under
the body by muscular action, and, being retained in this position
192 SPRING-TAILS CHAP,
of restraint by the catch, when the latter is removed the spring
extends by reason of its elasticity, and the leap is thus executed.
Whether this is really the exact method of leaping is, however,
doubtful, for Lubbock says that the catch “only exists in certain
genera”; while in its structure it does not appear to be well
calculated to retain in position an organ that by virtue of its
elasticity is constantly exerting a considerable force.
The ventral tube is an anomalous and enigmatic structure.
In the lower forms, such as Lipura or Anurida, it consists merely
of a papilla (Fig. 100, A, a) more or less divided by fissure into two
parts. In the Smynthuridae it is more highly developed, and
protects two long delicate tubes that are capable of being
protruded, as shown in the outline profile of Smynthurus fuscus
(Fig. 99), which is taken from specimens preserved in balsam by
Mr. J. J. Lister, The nature and use
of this ventral tube have given rise to
much discussion. Lubbock considered,
and others have agreed with him, that
it serves to attach the Insect to bodies
to which it may be desirable the Insect
should, when in the perpendicular posi-
tion, adhere. Reuter’ assigns a quite
different function to this singular struc-
Fic. 99.—Smynthurus fuscus, ture, He states that the hairs of the
with exsertile vesicle (a) pro- ;
truded from ventral tube; body are hygroscopic, and that the
0; the eprinp ogre a ces. peculiar claws of the Insect having
collected the moisture from the hairs, the ventral tube becomes
the means of introducing the liquid into the body. These Insects
possess, however, a mouth, and there seems to be no reason why
a complex apparatus should be required in addition to it for so
simple a purpose as the introduction of moisture to the interior of
the body. Haase finds? that Collembola can crawl on glass
without the aid of the ventral tube; he considers its function
to be physiological, and that it may probably be respiratory as it
has been suggested is the case with the vesicles of Thysanura.
The function of the ventral tube is certainly not yet satisfac-
torily elucidated. The vesicles contained in it are said to be
extruded by blood-pressure, and withdrawn by muscular action
in a manner similar to that which we have described as occurring
1 Ent. Tidskr. i. 1880, p. 159. 2 Morphol. Jahrb. xv. 1888, p. 361.
» Se sees s
VII APTERA 193
in the case of the exsertile vesicles of the Thysanura. The pro-
cesses in Smynthurus bear glandular structures at their ex-
tremities. It has been suggested that the ventral tube of Collem-
bola is the homologue of a pair of ventral appendages. The term
Collophore has been applied to it somewhat prematurely, seeing
the doubt that still exists as to its function.
‘Some of the Collembola possess a very curious structure
called the prostemmatic or ante-ocular organ; its nature and
function have been very inadequately investigated. The ocular
organs of the Collembola consist, when they are present, of
isolated ocelli placed at the sides of the head like the corre-
sponding organs of caterpillars; the prostemmate is placed
slightly in front of the group of ocelli, and has a concentric
arrangement of its parts, reminding one somewhat of the com-
pound eyes of the higher Insects. This structure is represented
in Fig. 100, B, C; it is said by Sir John Lubbock to be present
in some of the Lipuridae that have no ocelli, and he therefore
prefers to speak of it as the “ post-antennal ” organ. .
A very characteristic feature in the Collembola is the slight
development of the tracheal system. Although writers are far
from being in accord as to details, it seems that stigmata and
tracheae are usually absent. In Smynthurus there are, however,
according to Lubbock,—whose statement is confirmed by Meinert
and Tullberg,—a pair of stigmata situate on the head below the
antennae, and from these there extends a tracheal system through-
out the body. Such a position for stigmata is almost, if not
quite unique in Insects; Grassi, however, seems to have found
something of the kind existing in the embryo of the bee.
At present only a small number of species of the Order -
Aptera are known; Lubbock recognised about sixty British
species, and Finot sixty-five as found in France. The North
American forms’ have not received so much attention as the
European, and the Aptera of other countries, though they are
probably everywhere fairly numerous, are scarcely known at all. A
few have been described from the Indo-Malayan region and some
from Chili, and the writer has seen species from the West Indian
and Sandwich Islands. All the exotic forms as yet detected are
very similar to those of Europe.
The Thysanura are probably not very numerous in species, and
appear to be in general intolerant of cold. With the Collembola
VOL. V 0
194 APTERA CHAP.
the reverse is the case. They are excessively numerous in
individuals; they are found nearly everywhere on the surface of
the ground in climatic conditions like those of our country,
while no less than sixteen species have been found in Nova
Zembla and one each in Kerguelen and South Georgia. One
_ species, if not more, of Podura, lives on the surface of stagnant
waters, on which the minute creatures may frequently be seen
leaping about in great numbers after being disturbed.
In 1874 the plain of Gennevilliers in France was copiously
irrigated ; in the following year the soil was still very damp, and
there existed numerous pools of stagnant water, on the surface
of which Podura aquatica was developed in such prodigious
quantity as to excite the astonishment of the inhabitants of the
region. :
Accounts have been frequently given of the occurrence on
snow and glaciers of Insects spoken of as snow-fleas, or snow-
worms. These mostly relate to Poduridae, which are sometimes
found in countless number in such situations. The reason for
this is not well understood. According to F. Léw,' on the 17th
of March at St. Jacob in Carinthia, Parson Kaiser observed, on
the occurrence of the first thaw-weather, enormous numbers of a
‘Podura (? Achorutes murorum) on the surface of the snow for an
extent of about half a mile, the snow being rendered black in
appearance by them; eleven days afterwards they were found in
diminished numbers on the snow, but in large quantity on the |
water left by its melting. This account suggests that the
occurrence of the Insects on the snow was merely an incident
during their passage from the land, where they had been
hibernating, to the surface of the water.
One little member of the Lipuridae, Anurida maritima -
(Lipura maritima of Lubbock), has the habit, very unusual for an
Insect, of frequenting salt water. It lives amongst the rocks on —
the shores of the English Channel, between high and low tide-
marks. Its habits have been to some extent observed by
Laboulbéne ? and Moniez*; it appears to be gregarious, and when
the tide is high, to shelter itself against the commotions of the
water in chinks of the rocks and other positions of advan-
tage. When the tide is out the Insects apparently delight to
1 Verh. zool.-bot. Ges. Wien, viii. 1858, p. 564.
2 Ann. Soc. ent. France, 4th ser. iv. 1864, p. 705.
3 Rev. biol. Nord France, ii. 1890, p. 347.
/
@
fh
=
3 a
:
ot ea an Pee og Poet
completely covered with a coat
- touch it. The little creature
vie | APTERA 195
congregate in masses on the surface of the rock pools. This
Anurida can endure prolonged immersion; but both the ob-
servers we are quoting say that it is, when submerged, usually
of air so that the water does not
can, however, it would appear,
subsist for some time in the
pools of salt water, even when
it is not surrounded by its
customary protecting envelope
of the more congenial element.
Its food is said, on very slender
evidence, to consist of the re-
mains of small marine animals, 7,100, Anwida maritima: &, under
such as Molluscs. We repro- prostemmatic organ of young; C, of
duce some of Laboulbéne’s adult. (After Laboulbeéne, )
figures (Fig. 100); the under-surface shows at a the divided pap-
illa of the ventral tube ; B, C represent the peculiar prostemmatic
_ organ, alluded to on p. 193, in its mature and immature states.
Very little information exists as to the life-history of the
Aptera ; as for their food, it is generally considered to consist of
refuse vegetable or animal matter. It is usual to say that they
are completely destitute of metamorphosis, but Tenpleton says of
Lepisma niveo-fasciata that “the young differ so much from the
mature Insect that I took them at first for a distinct species; the
thoracic plates are proportionately less broad, and the first is
devoid of the white marginal band.” As regards the moults, it
would appear that in this, as in so many other points, great
diversity prevails, Grassi stating that in Campodea there is a
single fragmentary casting of the skin; and Sommer informing us
that in Macrotoma plumbea the moults are not only numerous,
but continue, after the creature has attained its full growth,
throughout life. |
A very marked feature of the Aptera is their intolerance of a
dry atmosphere. Although Campodea can exist under very
diverse conditions, it dies very soon after being placed in a dry
closed tube; and the same susceptibility appears to be shared by
all the other members of the Order, though it is not so extreme
in all; possibly it may be due to some peculiarity in the structure
196 APTERA CHAT.
of the integument. So far as tolerance of heat and cold goes,
the Aptera can apparently exist in. any climate, for though some
of the species extend to the Arctic regions, others are peculiar to
the tropics. |
Thysanura are recorded by Klebs and Scudder as occurring
commonly in amber; the latter author has described a fossil,
supposed to be a Lepisma, found in the Tertiary deposits at
Florissant. Scudder has also described another fossil, hkewise
from Florissant, which he considers to form a special sub-order of
Thysanura—Ballostoma—but it is extremely doubtful whether
this anomalous creature should be assigned to the Order at all. A
still older fossil, Dasyleptus lucasii. Brongniart, from the Carbon-
iferous strata in France, is considered to belong to the Order
Aptera, but it must be admitted there is some doubt on this
point.
The interest aroused in the minds of naturalists by the
comparatively simple forms of these purely wingless and therefore
anomalous Insects has been accompanied by much discussion as to
their relations to other Insects, and as to whether they are
really primitive forms, or whether they may perhaps be degenerate
descendants from some less unusual states of Insect-life. Mayer
and Brauer dissociated our Aptera entirely from other Insects,
and proposed to consider the Hexapoda as being composed of two
eroups—(1) the Apterygogenea, consisting of the few species we
have been specially considering ; and (2) the Pterygogenea, includ-
ing all the rest of the immense crowd of Insect forms. They
were not, however, able to accompany their proposed division by
any satisfactory characters of distinction, and the subsequent
‘progress of knowledge has not supported their view, all the best
investigators having found it necessary to recognise the extremely
intimate relations of these Insects with the Orthoptera. Meinert
thought that Lepisma must be included in the Orthoptera ; Grassi
proposes to consider the Thysanura as a distinct division of
Orthoptera ; and Oudemans recognises the close relations existing
between Machilis and Orthoptera proper. Finot includes the
Aptera in his Orthoptéres de la France, and a species of Japyx
has actually been described by a competent entomologist as an
apterous earwig. At present, therefore, we must conclude that
no good distinction has been found to justify the separation of .
“the Aptera from all other Insects.
VII APTERA 197
The taxonomy of the Collembola has not yet been adequately
treated, and it is possible that more grounds will be found
for separating them as a distinct Order from the Thysanura,—a
course that was advocated by Lubbock,—than exist for dividing
these latter from the Orthoptera proper. There are apparently no
grounds for considering the Aptera to be degenerate Insects, and we
may adopt the view of Grassi, that they are primitive, or rather
little evolved forms. It must be admitted that there are not at
‘present any sufficient reasons for considering these Insects to be
“ancient ” or “ancestral.” The vague general resemblance of Cam-
podea to many young Insects of very different kinds is clearly the
correlative of its simple form, and is no more proof of actual
‘ancestry to them than their resemblances inter se are proofs of
ancestry to one another. But even if deprived of its claim to
antiquity and to ancestral honours, it must be admitted that
Campodea is an interesting creature. In its structure one of the
most fragile of organisms, with a very feeble respiratory system,
inadequate organs of sense, only one pair of ovarian tubes, very
imperfect mouth-organs, and a simple alimentary canal, it
nevertheless flourishes while highly-endowed Insects become
extinct. In the suburban gardens of London, on the shores of —
the Mediterranean, on the summits of the higher Pyrenees, in
North America even it is said in the caves of Kentucky, and
in India, Campodea is at home, and will probably always be
with us.
CHAPTER VIII
ORTHOPTERA——-FORFICULIDAE, EARWIGS——HEMIMERIDAE
Order II.—Orthoptera.
Insects with the mouth parts conspicuous, formed for biting, the
Sour palpi very distinct, the lower lip longitudinally divided
in the middle. The tegmina (mesothoracie wings), of parch-
ment-like consistence, in repose closed on the back of the
Insect so as to protect it. The metathoracie wings, of more
delicate consistence, ample, furnished with radiating or
divergent nervures starting from the point of articulation,
and with short cross nervules forming a sort of network;
in repose collapsing like a fan, and more or less completely
covered by the tegmina (except in certain Phasmidae, where,
though the wings are ample, the tegmina are minute, so that
the wings are uncovered). Ina few forms (winged Forfi-
culidae and some Blattidae) the metathoracie wings are, in
addition to the longitudinal folding, contracted by means of
one or two transverse folds. ' The mode of growth of each
individual is a gradual increase of size, without any abrupt
change of form, except that the wings are only fully developed in
the final condition. There is no special pupal instar. Species
in which the wings are absent or rudimentary are numerous.
THE Orthoptera are Insects of comparatively large size. The
Order, indeed, includes the largest of existing Insects, while none |
are so minute as many of the members of the other Orders are ;
three millimetres is the least length known for an Orthopterous
Insect, and there are very few so small, though this is ten times
the length of the smallest beetle. The Order includes earwigs,
cockroaches, soothsayers or praying-insects, stick- and leaf-insects,
grasshoppers, locusts, green grasshoppers, and crickets.
CHAP. VIII ORTHOPTERA 199
The changes of form that accompany the growth of the
individual are much less abrupt and conspicuous than they are
in most other Insects. The metamorphosis is therefore called
Paurometabolous. It has been supposed by some naturalists that
Orthoptera go through a larger portion of their development in
the egg than other Insects do. This does not clearly appear -
to be the case, though. it seems that there are distinctions of a
general character in the embryology; the period of development
in the egg is prolonged, and the yolk is said by Wheeler? to be
more than usually abundant in comparison with the size of the
young embryo. The embryonic development may in tropical
countries be accomplished in three weeks (see Mantidae), but in
countries where winter supervenes, the period may in some
species be extended over seven or eight months.
The external features of the post-embryonic development—a
term that is more convenient in connexion with Orthoptera than
metamorphosis —are as follows: the wings are never present when
the Insect is first hatched, but appear subsequently, and increase
in size at the moults; the form and proportions of the segments
of the Bidy especially of the thorax—undergo much change ;
an alteration of colour occurs at some of the moults, and the
integument becomes harder in the adult condition. Neither
the development of the internal organs, nor the physiological pro-
cesses by which the changes of external form are effected, appear
to have been studied to any great extent.
Many of the Orthoptera do not possess wings fit for flight, and
some species even in the adult state have no trace whatever of such
organs. Flight, indeed, appears to be of minor importance in the
Order; in many cases where the wings exist they are purely musical
organs, and are not of any use for flight. The apterous and the
flightless conditions are not confined to one division of the Order,
but are found in all the families and in many of their sub-
divisions. As the front pair of wings in Orthoptera do not really
carry out the function of flight, and as they differ in several par-
ticulars from the hinder pair, or true wings, it is usual to call
them tegmina.. The musical powers of the Orthoptera are confined
to the saltatorial group of families. The Cursoria are dumb or
nearly so; it is a remarkable fact that also in this latter division
the alar organs, though frequently present, have but little value
1 J. Morphol. viii. 1893, p. 64.
200 . ORTHOPTERA CHAP.
for flight, and are in some cases devoted to what we may call
purposes of ornament or concealment. This is specially the case
in the Phasmidae and Mantidae, where the effectiveness of colour
and pattern of these parts becomes truly astonishing. The
tegmina frequently exhibit an extraordinary resemblance to
vegetable structures, and this appearance is not superficial, for it —
may be seen that the nervures of the wings in their disposition and
appearance resemble almost exactly the ribs of leaves. One of the
most remarkable of the features of Orthoptera is that a great
difference frequently exists between the colours of the tegmina
and of the wings, ze. the front and hind wings; the latter are
concealed in the condition of repose, but when activity is entered
on and they are displayed, the individual becomes in appearance
a totally different creature. In some cases, contrary to what
usually occurs in Insects, it is the female that is most remark-
able; the male in Mantidae and Phasmidae being frequently a
creature of quite inferior appearance and power in comparison
with his consort. The musical powers of the saltatorial
Orthoptera are, however, specially characteristic of the male sex.
There is evidence that these powers are of great importance to
the creatures, though in what way is far from clear. Some parts
of the structures of the body are in many of these musical species
clearly dominated by the musical organs, and are apparently
specially directed to
securing their effici-
ency. We find in some
Locustidae that the
tegmina are nothing
but sound - producing
instruments, while the
pronotum is prolonged
Fic. 101.—Poecilimon afinis 6. Bulgaria. Alar organs to form a hood that
serving only as musical organs. The ear on front protects them without
tibia and aural orifice of prothorax are well shown. ‘ °
encumbering their ac-
tion. In the males of the Pneumorides, where the phonetic organ
is situated on the abdomen, this part of the body is inflated and
tense, no doubt with the result of increasing the volume and quality
of the sound. In the genus Methone (Fig. 185) we find a grass-
hopper whose great hind legs have no saltatorial function, and but
little power of locomotion, but act as parts of a sound-producing
vil 7 THE FAMILIES , 201
instrument, and as agents for protecting some parts of the body
in repose. Further particulars of these cases must be looked for
in our accounts of the different groups.
The eggs of many Orthoptera are deposited in capsules or
cases; these capsules may contain only one egg, or a great many.
The Order includes many species of Insects, though in Britain
it is poorly represented: we have only about forty species, and
this small number includes some that are naturalised. Only a
few of the forty extend their range to Scotland. A revision of
the species found in Britain has recently been made by Mr.
Eland Shaw.’ In continental Europe, especially in the south,
the species become more numerous; about 500 are known as
inhabitants of geographical Europe. In countries where the face
of nature has been less transformed by the operations of man,
’ and especially in the tropical parts of the world, Orthoptera are.
much more abundant.
The lowest number at which the species now existing on the
surface of the earth can be estimated is 10,000. This, however,
is probably far under the mark, for the smaller and more obscure
species of Orthoptera have never been thoroughly collected in
any tropical continental region, while new forms of even the
largest size are still frequently discovered in the tropics.
We shall treat the Order as composed of eight families :—
1. Forficulidae—Tegmina short, wings complexly folded ; body
armed at the extremity with strong forceps.
Series, Cursoria ; |2. Hemimeridae—Apterous: head exserted, constricted behind.
hind legs but | 3. Blattidae—Coxae of the legs large, exserted, protecting
little different the lower part of the body.
from the others. | 4. Mantidae—Front legs very large, raptorial, armed with spines.
5. Phasmidae—Mesothorax ec as compared with the pro-
thorax.
Series, Saltatoria : | 6. Acridiidae—Antennae short, not setaceous, of not more than
hind legs elon- 30 joints, tarsi three-jointed.
gate, formed for } 7. Locustidae—Antennae very long, setaceous, composed of a
leaping, their large number of joints, tarsi four-jointed.
femora usually | g
thickened.
“I
. Gryllidae—Antennae very long, setaceous, tarsi two- or three-
jointed.
The first five of these subdivisions are amongst the. most
distinct of any that exist in the Insecta, there being no con-
necting links between them. The three groups forming the
1 Ent. Mo. Mag. xxv. 1889, and xxvi. 1890.
202 ORTHOPTERA CHAP.
Saltatoria are much more intimately allied, and should, taken
together, probably have only the same taxonomic value as any one
of the other five groups. |
Owing partly to the inherent difficulties of the subject, and
partly to the fragmentary manner in which it has been treated
by systematists, it has been impossible till recently to form any
clear idea of the classification of Orthoptera. During the last
twenty years Henri de Saussure and Brunner von Wattenwyl
have greatly elucidated this subject. The latter of these two
distinguished naturalists has recently published} a revision of the
system of Orthoptera, which will be of great assistance to those
who may wish to study these Insects. We therefore reproduce
from it the characters of the tribes, placing the portion relating
to each family at the end of our sketch thereof.
Fam. I. Forficulidae—Earwigs.
(DERMAPTERA OR DERMATOPTERA OF BRAUER AND OTHERS)
Insects of elongate form, with an imbricate arrangement of the
segments of the body; bearing at
the posterior extremity a pair of
callipers or more distorted instru-
ments. The hind wings (when
present) folded in a complea
manner, and covered, except at their
tips, by a pair of short wing-covers
(tegmina), of a leather-like consist-
ence. Wingless forms are very
numerous. The young is very
similar to the adult.
- Although earwigs are said to be
rare in most parts of the world, yet
| in Europe no Insect is better known
Fic. 102.—Pygidicrana hugeli. than Forficula auricularia, the common
qin earwig, it being very abundant even in
gardens and cultivated places. In certain seasons it not un-
frequently enters our houses, in which case it too often falls a _
1 Ann. Mus. Genova, xxxiii. (1892).
VIL EARWIGS 203
victim to prejudices that have very little to justify them. This
Insect is a good type of the winged earwigs. In the parts of
the mouth it exhibits the structures usual in the Orthoptera ;
there is a large labrum, a pair of maxillae, each provided with two
lobes and a palpus consisting of two very short basal joints and
three longer joints beyond these ; the mandibles are strong, with
curvate pointed extremities; in the lower lip there is a ligula
exposed in front of a very large mentum; it consists of two
pieces, not joined together along the middle, but each bearing on
its lateral edge a palpus with two elongate joints and a short
basal one; this lip is completed by the lingua, which reposes
on the upper face of the part, and completely overlaps and
protects the chink left by the want of union along the middle
line of the external parts of the lip. The antennae are elon-
gate, filiform, and are borne very near the front of the exserted
head. There are rather large facetted eyes, but no ocelli. The
three segments of the thorax are distinct, the prothorax being
quite free and capable of movement independent of the parts
behind it: the meso- and meta-nota are covered by the tegmina
and wings; these latter project slightly from underneath the
former in the shape of small slips, that are often of rather lighter
colour; the wing-covers are short, not extending beyond the
insertion of the hind legs, and repose flat on the back, meeting
together in a straight line along the middle. These peculiar
flat, abbreviated wing-covers, with small slips (which are portions
. of the folded wings) ‘projecting a little from underneath them,
are distinctive marks of the winged Forficulidae.
The legs are inserted far from one another, the coxae being
small; each sternum of the three thoracic segments projects
backwards, forming a peculiar long, free fold, underlapping the
front part of the following segment. The hind body or abdomen
is elongate, and is. formed of ten segments; the number readily
visible being two less in the female than it is in the male.
The segments are fitted together by a complex imbrication,
which admits of great mobility and distension, while offering a
remarkable power of resistance to external pressure: each
segment is inserted far forward in the interior of that preceding
it, and each also consists of separate upper and lower plates that
much overlap where they meet at the sides (see Fig. 103). The
body is always terminated by a pair of horny, pincer-like
204 ORTHOPTERA CHAP.
processes, which are differently shaped according to the sex of
the individual.
The structure of the abdomen in the earwig has given rise to
considerable discussion. In Fig. 103 we reproduce Westwood’s
diagram of it as seen fully distended
in a female specimen; in this state the -
minute spiracles can be detected, though
in the normal condition of the body
they cannot be seen, being placed on
the delicate membranes that connect
the chitinous plates. Westwood’s inter-
pretation of the structure was not, how-
ever, quite correct, as the part which .
he considered to be the first dorsal
plate is really the second; so that the
Frc. 103.—Lateral view of For. Segments numbered 7, 8, 9 in our
sie uenaad snomng’ Sure are really 8,9, 10. ‘The com
spiracles, S$, and the small Mon earwig 18 interesting as exhibit-
Sth and 9th dorsal plates ing in an imperfect state, the union
(7 and 8 in Fig.).
of the first dorsal plate of the abdo-
men with the thorax; a condition which is carried to so
great an extent in the Hymenoptera as to quite obscure the
nature of the parts, and which has consequently given rise to
much perplexity and discussion. We repre-
sent this structure as seen in the common
earwig in Fig. 104, where a represents the
pronotum, ) the mesonotum, ¢ the metanotum,
d the first, f the second abdominal segment ;
e being a delicate membrane of considerable
size that intervenes between the two, and
which is more exposed than are the corre-
sponding membranes connecting the subse- yy, 104,—Dorsal por-
quent rings ; a condition similar to that which tions of the middle
: ‘ ; segments of body of
is found in Cimbex, Cephus, and some other = Forficula auricul- -
Hymenoptera. — aria (tegmina and
wings removed).
On the under surface of the abdomen of the
earwig the full number of 10 plates cannot be superficially dis-
tinguished ; but it is found by dissection that in the female the
short eighth and ninth dorsal rings are joined on the ventral aspect
by a delicate membrane, while the tenth ventral is of a less delicate
VIII . WINGLESS EARWIGS 205
nature, and forms a triangular plate at the base of each half of
the forceps. Between the branches of the forceps there is a per-
pendicular plate, the pygidium of Orthopterists, possibly the
unpaired terminal portion of the body seen in some embryos, and -
called the telson. The pygidium is a separate sclerite, though
it looks as if it were only a portion of the large tenth dessa
plate bent downwards, and in some descriptive works is errone-
ously described as being such.
A very large number of species of Forficulidae have the
organs of flight undeveloped. Fig. 105 represents Chelidura
dilatata, an apterous form that is very com-
mon in the Eastern Pyrenees. The condition
of the meso- and meta-nota—the parts from
which the tegmina and wings are developed,
and to which they are attached when present
—is very remarkable in these forms, and
exhibits much variety. In Fig. 106 we
represent the conditions of these parts in
a few apterous forms. The tegmina or the
segment from which they are developed (0),
are seen in the shape of a plate which may
extend all across the middle and be undi-
vided (No. 4); in which case the appearance indicates entire
absence of the tegmina; these are, on the contrary, evidently present
in the form of slips grafted one to each side of the second thoracic
seoment in Anisolabis (No. 3); or they may look like. short
| broad slips extending all
pee - across the body, and mark-
i) 1 Janes ing off a piece frequently
c = calledascutellum, but which
£ Wo Le is really the mesonotum
a.
(some species of Chelidura,
Fic. 106. —Tegmina and wings cently in part or gg No, 2): ars again, they
invisible) of apterous earwigs. 1, Chelidura sp.;
2, Chelidura dilatata ; 3, Anisolabis moesta.; ae be nearly fr ee teg-
EAcgrerina, 4 Bist tomate ements?» anina, somewhat, similar to
"those of the winged forms ;
this is the case with some species of Chelidura, as represented
by No.1. This last figure is taken from a species from the
Sierra Nevada, apparently undescribed, allied to C. bolivari.
In the cases we are considering no analogous structures exist on
Fic. 105.—Chelidura
dilatata, male. Pyrenees.
206 ORTHOPTERA CHAP.
the metanotum (the part of the body that in the winged forms
bears the wings, and which is marked ¢ in our diagrams, Fig.
106), so that the tegmina are to all appearance less rudimentary ~
(or vestigial) than the wings. The metanotum forms a sort of
flap, called by Fischer? “involucrum alarum”; he considered
the part ‘immediately behind this to be the metanotum; this —
piece is, however, no doubt really part of the abdomen (d in our
Figure). This is apparently the view taken by Brunner.” The ~
structure of these parts is important as bearing on the subject
of the nature and origin of Insects’ wings, a question to which
no satisfactory answer has yet been given. The appearances
we have remarked on are to some extent similar to the con-
ditions existing in the immature state of the organs of flight in
the common earwig (see Fig. 112, p. 212), but whether the
varieties presented by the wingless forms have parallels in the
immature conditions of the various winged forms is quite
uncertain, the life-histories of earwigs being almost unknown.
The developed wings of earwigs are worthy of attention,
both as regards their actual structure and the manner in which
they are folded up in repose. When
expanded they have a shape curiously
suggestive of the human ear. The chief
parts of the wing, as shown in Fig. 107,
A, are a, b, two portions of the horny
piece that forms the scale which covers
the more delicate parts of the wing
when it is folded, and which, according
fia. 1b oes ee ee to Brunner, represents the radial and
auricularia, A, Wing ex- Ulnar fields of the wings of Acridiidae
gern Se ipa iy berks and Locustidae (see Fig. 167); ¢€ is
‘the small apical field limited below
by the vena dividens; d is the vena plicata which runs
along the under side of the scale as far as the apical field,
where it gives off the axillary nerves; e is a vena spuria, or
adventitious vein such as exists in many other Orthoptera
with delicate wings. On the front part of the scale, a, and
on a different plane so that it is not shown in our figure,
there is a very delicate small band which is supposed to repre-
1 Orthoptera Europaea, 1853, pl. vi. f. 4, p. 484.
2 Morph. Bedeut. Seg. Orthopt. 1876, p. 14; and Prod. Orthopt. Europ. 1882, p. 3.
VIII WINGS OF EARWIGS 207
sent the marginal field of the wing of other Orthoptera. There
are, however, grave difficulties in the way of accepting this view
of the earwig’s wing, amongst which we may mention the
position of the vena dividens and its relation to the so-called
radial and ulnar fields of the wing. The wings are remarkable
for their delicacy ; moreover, the way in which they fold up so
as to be packed in the manner shown in B, Fig. 107, is very
interesting, there being, in fact, no other Insects that fold up
their wings in so complicated and compact a fashion as the
earwigs do. The process is carried out somewhat as follows: the
longer radii come a little nearer together, the delicate membrane
between them falling into folds somewhat like those of a paper
fan; a- transverse fold, or turn-over, then occurs at the point
where the radii, or axillary nerves, start from the vena plicata ;
then a second transverse fold, but in a reversed direction, occurs
affecting the wing just close to the spots where the shorter
radial nervures are dilated; then by a contraction close to the
scale the whole series of complex folds and double are brought
together and compressed. ,
It is quite a mystery why earwigs should fold their wings
in this complex manner, and it is still more remarkable that
the Insects very rarely use them. Indeed, though orjicula
auricularia is scarcely surpassed in numbers by any British
Insect, yet it is rarely seen on the wing; it is probable that
the majority of the individuals of this species may never make
use of their organs of flight or go through the complex process
of unfolding and folding them. It should be remarked that no
part of the delicate membranous expanse of the wing is exposed
when the wings are packed in their position of repose; for the
portion that projects from under the tegmina—and which, it
will be remembered, is always present, for when wings exist
in earwigs they are never entirely concealed by the tegmina—
is, it is curious to note, of hard texture, and is frequently coloured
and sculptured in harmony with the tegmen; in fact, one small
part of the wing forms in colour and texture a most striking
contrast to the rest of the organ, but agrees in these respects
with the wing-covers. This condition is seen in Fig. 108,
where B shows the sculpture of the tegmina #, and of the
projecting tips of the wings w. There are numerous other
instances in Orthoptera where one part of a wing or wing-case
208 ORTHOPTERA CHAP.
is exposed and the other part concealed, and the exposed portion
is totally different in colour and texture from the concealed
portion.
The wings of earwigs are attached to the body in a very
unusual manner ;.each wing is continued
inwards on the upper surface of the
metanotum, as if it were a layer of the
integument meeting its fellow on the
‘mesial line; the point of contact forming
two angles scat behind the metanotum.
Some writers have considered that the
tegmina of earwigs are not the homologues
of those of other Orthoptera, but are really
tegulae (cf. Fig. 56, p. 103). We are not
aware that any direct evidence has been
produced in support of this view.
The pair of forceps with which the
body is armed at its extremity forms
another character almost peculiar to the
earwigs, but which exists in the genus
Japyx of the Thysanura. These forceps
> ge age eee syiae vary much in the different genera of the
line of the Insect; B, family ; they sometimes attain a large size
vies eh : shoving th and assume very extraordinary and dis-
similar sculpture, _ torted shapes. They are occasionally used
by the Insects as a means of completing the process of packing
up the wings, but in many species it is not probable that they
can be used for this purpose, because their great size and peculiarly
‘distorted forms render them unsuitable for assisting in a delicate
process of arrangement ; they are, too, always present in the wingless
forms of the family. Their importance to the creature is at
present quite obscure; we can only compare them with the
horns of Lamellicorn Coleoptera, which have hitherto proved
inexplicable so far as utility is concerned. No doubt the
callipers of the earwigs give them an imposing appearance, and
may be of some little advantage on this account; they are not
known to be used as offensive instruments for fighting, but they
are occasionally brought into play for purposes of defence, the
creatures using them for the infliction of nips, which, however,
are by no means of a formidable character.
:
'
‘
a
=
Seal
Vul EARWIG-FORCEPS 209
These forceps are, in the case of the common earwig—and they
have not been studied from this point of view in any other
species—remarkable, because of the great variation in their
development in the male, a character which again reminds us
of the horns of Lamellicorn beetles: in the female they are
comparatively invariable, as is also the
case in the few species of Lamellicornia,
which possess horned females. A and :
B in Figure 109 represent the forceps
A B c
of different males of the common earwig,
C showing those of the other sex. The
subject of the variation of the male
callipers of the earwig has been con-
sidered by Messrs. Bateson and Brind- F109. — Forceps of the
. : : common earwig: A, of large
ley who examined .1000 specimens male; B, of small male;
captured on the same day on one of the ©&% ° female.
Farne islands off the coast of Northumberland ; 583 of these were
mature males, and the pincers were found to vary in length
from about 24 mm. to 9 mm. (A and B in Fig. 109 repre-
sent two of the more extreme forms of this set of individuals.)
Specimens of medium size were not, as it might perhaps have
been expected they would be, the most common; there were,
in fact, only about 12 individuals having the forceps of the
medium length—4#? to 54 mm., while there were no less than
90 individuals having forceps of a length of about 7 mm., and
120 with a length of from 2? to 34. Males with a medium
large length of the organ and with a medium small length
thereof were the most abundant, so that a sort of dimorphism
was found to exist. Similar relations were detected in the
length of the horns of the male of a Lamellicorn beetle examined
by these gentlemen. In the case of the set of earwigs we have
mentioned, very little variation existed in the length of the
forceps in the female sex.
In many earwigs—including F. auricularia—there may be
seen on each side of the dorsal aspect of the true fourth, or of the
fourth and neighbouring segments of the hind body a small
elevation, called by systematists a plica or fold, and on examina-
tion the fold will be found to possess a small orifice on its
posterior aspect. These folds are shown in Figs. 105 and 108;
1 Proc. Zool. Soc. London, 1892, p. 586,
VOL. V P
210 ORTHOPTERA CHAP.
they have been made use of for purposes of classification, though
no functional importance was attached to them. Meinert,
however, discovered’ that there are foetid glands in this
situation, and Vosseler has recently shown? that the folds are
connected with scent-glands, from which proceed, in all pro-
bability, the peculiar odour that is sometimes given off by the
earwig. The forms destitute of the folds, eg. Labidura, are
considered to have no scent glands. There is a very peculiar
series of smooth marks in the earwigs on the dorsal aspect of
the abdominal segments, and these are present in the glandless
forms.as well as in the others.
The internal anatomy has been to some extent investigated
by Dufour and Meinert. Dufour dis-
/ sected F. auricularia and Labidura riparia,
and found® that salivary glands exist in
the latter Insect (called by him Forficula
gigantea), though he was unable to discover
them in the common earwig. According
to Meinert,* there are, however, salivary
glands affixed to the stipes of the maxillae
in / auricularia, while (in addition?) JL.
riparia possesses very elongate glands seated
in the middle or posterior part of the breast.
The alimentary canal is destitute of con-
volutions, but oesophagus, crop, and gizzard
all exist, and the intestine behind the
stomach consists of three divisions. The
Malpighian tubes are numerous, 30 or 40,
and elongate. The respiratory system is
not highly developed. Earwigs—the Euro-
pean species at least—have, as already mentioned, very small
powers of flight; the tracheal system is correspondingly small,
and is destitute of the vesicular dilatations that are so remarkable
in the migratory Locusts.
The three thoracic spiracles’ are readily observed in living
>)
Le,
oy, . F
Yl
=a
AT
— |
Lh ~~
ur)
é =
a
“
Whiz
=
> > EE Be
Fig. 110.—Labidura ripa-
_ gta, male. Europe.
1 Naturhistorisk Tidsskrift, 3rd ser. ii. 1868, p. 475.
2 Arch. mikr. Anat. xxxvi. 1890, p. 565.
3 Ann. Sci. Nat. xiii. 1828, p. 337.
* Naturhistorisk Tidsskrift, 8rd ser. ii. 1868, p. 475.
° Some writers are of opinion that there are only two thoracic spiracles in Insects,
considering the third as belonging really to the abdomen. Looking on the point as
ae a ae Se
¥
obliquely directed lateral part of the
“yII EARWIGS 211
individuals. There are seven pairs of abdominal spiracles, which,
however, are very minute, and can only be found by distending
the body as shown in Fig. 103. The ventral chain consists of
nine ganglia (the sub-oesophageal centre is not alluded to by
Dufour); the three thoracic are equidistant and rather small;
the hindmost of the six abdominal ganglia is considerably larger
than any one of the other five.
The ovaries of Labidura riparia and Forficula auricularia are
extremely different. In Z. riparia there are on each side five
tubes, each terminating separately in an
oviduct. In F& auricularia there is but
one tube on each side, but it is covered
by three longitudinal series of very short
sub-sessile, grape-like bodies, each of the
two tubes being much dilated behind the
point where these bodies cease.
The testes in earwigs are peculiar and
simple; they consist, on each side, of a
pair of curvate tubular bodies, connected
at their bases and prolonged outwards in
the form of an elongate, slender vas de-
ferens. The structures in the males of
several species have been described at
some length by Meinert,’ who finds that
in some species a double ejaculatory duct
exists.
The young is similar to the adult in :
form; in the winged forms it is always pyc, 111.—Ovaries of Labi-
easy to distinguish the adult by the full @ureriparia, A; and For-
; ‘ jficula auricularia, B.
development of the wings, but in the ‘(After Dufour.)
Wingless forms it can only be decided
with certainty that a specimen is not adult by the softer and
weaker condition of the integuments. Scarcely anything appears
to be known as to the life-history, except a few observa-
tions that have been made on the common earwig; Camerano
found? that this Insect has certainly three ecdyses, and possibly
at present chiefly one of nomenclature, we make use of the more usual mode of
expression.
1 As on last page, and also op. cit. v. 1868, p. 278.
2 Bull. Ent. Ital. xii. 1880, p. 46.
212 ORTHOPTERA CHAP.
an earlier one which he failed to notice, and his observa-
tion confirms the vague previous statement of Fischer. The
egos, in the neighbourhood of Turin, are deposited and hatched
in the early spring; in one case they were laid on the 10th
March, and the Insects issuing from them had completed their
growth and were transformed into perfect Insects on the 22nd.
May. In the immature state the
alar structures of the future imago
may be detected. The tegmina-bear-
ing sclerites, ¢, Fig. 112, look then
somewhat like those of some of the
apterous forms (Fig. 106) and, as
shown in A and B, Fig. 112, do not
Fic. 112.—Notal plates from which differ greatly in the earlier and later
arp enna wr stages. The wings, however, change
young, A, and more advanced, B, much more than the tegmina do;
nymph. at first (Fig. 112, A) there is
but little difference between the two, though in the interior of
the wing-flap some traces of a radiate arrangement can be seen,
as shown at W in A, Fig. 112; in a subsequent condition the
wing-pads are increased in. size and are more divided, the appear-
ance indicating that the wings themselves are present and packed
about a centre, as shown in W of B, Fig. 112.
In the young of the common earwig the number of joints? in
the antennae increases with age. Camerano, l.c., says that before -
emergence from the egg there are apparently only 8 joints in the
antennae, and Fischer states that the larvae of -#! auricularia
have at first only 8 antennal joints; later on 12 joints are
commonly found, and, according to Bateson? this number
occasionally persists even in the adult individual. Meinert says *
that the newly hatched Forfiewla has either 6 or 8 joints, and
he adds that in the later portion of the preparatory stage the
number is 12. Considerable discrepancy prevails in books as. to
the normal number of joints in the antennae of the adult /
auricularia, the statements varying from 13 to 15. The latter
number may be set aside as erroneous, although it is, curiously
1 It may be worth while to repeat that “ geal means a piece, and is the
equivalent of ‘‘link” in a chain.
2 Materials for the Study of Variation, 1894, p. 413.
> Naturhist. Tidsskrift, 3rd ser. ii. 1863, p. 474.
LI EE ec RT Lt .148.—Loncho-
really such by Westwood, who describes the abdo- MUAer deci,
men as consisting of nine segments. The flat apical _Pelago. (Alter
; estwood. )
appendages are attached behind the tenth dorsal
plate. The ventral plates are similar to the dorsal in arrangement,
except that in the female the eighth plate forms a sort of spoon-like
or gutter-like process to assist in carrying or depositing the eggs, and
that the two following segments are concealed by it, and are some-
times of more delicate texture. The legs vary greatly in the details of
Ni-
262 ORTHOPTERA > CHAP,
their shape: the coxae are short, oval, or round, never .large; the
trochanter is small; the front femora.
often have the basal part narrower
than the apical, and they are fre-
quently so formed that they can be
stretched out in front of the head, —
concealing its sides and outline and.
entirely encasing the antennae.
There is an arolium or cushion
between the claws of the five-
jointed tarsi. The front legs are
frequently longer than the others.
Only a very slight study has been
made of the alar organs of Phas-
midae; but according to Redten-
bacher and Brauer, they differ
greatly from those of Blattidae and
Mantidae, inasmuch as the costal
vein is placed not on the actual
i margin of the wing but in the
‘ field thereof, and in this respect
they more resemble the Orthoptera
saltatoria.
Very little information exists
as to the internal anatomy of the Phasmidae. Many years ago
a memoir of a fragmentary and discursive nature was published
on the subject by J. Miiller,’ but his conclusions require con-
firmation ; the nervous system, according to his account, which
refers to Arumatia ferula, has the anterior ganglia small, the
supra-oesophageal ganglion being apparently not larger than
those forming the ventral chain.
Joly’s more recent memoir on the anatomy of Phylliwm
crurifolium ? is also meagre; he states that the nervous system
resembles that of the locusts (Acridiidae), though there are at
least ten pairs of ganglia—one supra-, one infra-oesophageal, three
thoracic, and five abdominal. He found no salivary glands; the
Malpighian tubules are slender, elongate, and very numerous.
The tracheal system has no air-vesicles. He found no distinction
Fic. 149.—Heteropteryx grayi, male.
‘Borneo. One-half natural size.
1 Acta Ac. German. xii. 1825, pp. 555-672, pls. 1.-liv.
2 Mem. Ac. Sci. Toulouse, series 7, iii. pp. 1-30.
se
xr PHASMIDAE 263
of crop and proventriculus, but the true stomach appears to consist
of two different parts, the anterior being remarkably uneven
externally, though destitute of coeca, while on the posterior part
there are peculiar vermiform pronterss There are eighteen or
twenty tubes in each ovary.
When the young Insect is in the egg, ready for emergence,
the meso- and meta-thorax are not remarkably elongate, so that
the femora are not very far
apart, but by the time the crea-
ture has fairly emerged from the
prison of its embryonic life the
thoracic segments have attained
their usual proportions; much
expansion of the body takes place
as the Insect leaves the egg, so.
that it appears a marvel how it
could have been contained therein;
this expansion affects the parts
of the body unequally.
_ The records as to the post-
embryonic development of Phas-
midae are very scanty, but indi-
cate great differences in the
length of time occupied by it.
Bacillus patellifer is said to
moult several times, Diaphero-
mera femorata only twice. This
latter > ee becomes full iprtaies Fia.150.—Aschipasma catadromus, female.
in six weeks, while, according Sumatra. Natural size. (After West-
to Murray,’ Phylliwm scythe i in
required fifteen or sixteen months for growth, and did not
moult until ten months after hatching; the number of
ecdyses in the case of the Phylliwum was three. At cach
ehange of skin an immediate increase in.size, similar to that we
have noticed as occurring on leaving the egg, takes place; each
limb on being freed becoming about a fourth longer and larger
than the corresponding part of the envelope from which it has
just been withdrawn. After the second moult of Phyllium the
tegmina and wings made their appearance, but remained of very
1 Edinburgh Philosoph. Journ. January 1856.
264 ORTHOPTERA 7 CHAP.
small size until after the third moult, when they suddenly shot out
to their full size; they came out of little cases about a quarter of
an inch long, and in the course of a
few minutes attained their full size
of about two and a half inches of
length. In the apterous species the
difference between the young and
adults in external characters is very
slight. — ;
Phasmidae are very sensitive to
cold; both in North America and
Australia their lives are terminated —
by the occurrence of frost. They
are all vegetable feeders, the canni-
balism that has been attributed to
them by several writers being prob-
ably imaginary. They are, how-
ever, excessively voracious, so that a
pair will destroy a great quantity
of foliage; they are consequently
in some parts of the world classed
amongst injurious Insects. In Fiji
tT _ and the Friendly Islands, Lopaphus
Fie. 151.—Ceroys saevissima. Brazil. .
(After Westwood. ) cocophagus eats the cocoa-nut foliage
and causes a scarcity of food, so that —
it becomes a matter of necessity to destroy these Insects, One
writer has gone so far as to attribute the occurrence of cannibal
habits amongst the inhabitants of some of these islands to the want
of food caused by the ravages of this Insect. Some, if not all, of
the Phasmidae have the habit of ejecting a stinking fluid, that is
said to be very acrid, and occasionally, when it strikes the eye, to
cause blindness; this liquid comes from glands placed in the
thorax. Some Phasmidae are much relished as food by birds;
Napheromera femorata is sucked by several bugs as well as eaten
by birds, and another species is recorded to have harboured
Ichneumon-flies in its body without suffering any apparent incon-
venience from their presence or from their emergence. Not-
withstanding the great amount of food they consume and their
want of activity, they produce comparatively few eggs. From
twelve to twenty or thirty is frequently mentioned as about the
XI EGGS OF PHASMIDAE 265
number, but in the case of Diapheromera femorata Riley speaks
of upwards of one hundred. These eggs are not deposited in any
careful way, but are discharged at random, simply dropping from
the female; the noise caused by the dropping of the eggs of
MNapheromera femorata from the trees on which the Insects are
feeding to the ground is said to resemble the pattering of rain-
drops. The eggs of this
species-often remain till
the second year before
they hatch. The eggs
in the Phasmidae gen-
erally are of a most
remarkable nature, and
nearly every one who
; h k A B C D
mentions them Speaks Pye, 152.—Eggs of Phasmidae: A, Lonchodes duiven-
of their extreme resem- bodi ; B, Platycrania edulis ; C, Haplopus grayi ;
blance to seeds. Gdoldi2 D, Phyllium siccifolium. (After Kaup.)
has suggested that this is for the purpose of deceiving Ichneumons ;
it 1s, however, on record that the eggs are actually destroyed by
Ichneumons. It is worthy of notice that the eggs are-shed like
seeds, being dropped loosely and, as we have said, remaining on
the ground or elsewhere, sometimes for nearly two years, without
other protection than that they derive from their coverings.
Each egg is really a capsule containing an egg, reminding us thus
of the capsule of the Blattidae, which contains, however, always
a number of eggs. Not only do the eggs have a history like that
of seeds, and resemble them in appearance, but their capsule in
minute structure, as we shall subsequently show, greatly resembles
vegetable tissue. The egg-capsule in Phasmidae is provided
with a lid, which is pushed off when the Insect emerges (Fig.
157). This capsule induced Murray to suppose that the egg
contained within is really a pupa, and he argued therefrom that
in the Orthoptera the larval stages are passed in the egg, and
that the Insect after its emergence should be looked on as an
active pupa that takes food.
The individuals of this group of Insects possess the power of
reproducing a lost hmb; and Scudder, who has made some experi-
ments as to this,” states that if a leg be cut off beyond the
1 Zool. Jahrb. Syst. i. 1886, p. 724.
2 P. Boston Soc. xii. 1869, p. 99.
266 - ORTHOPTERA CHAP.
trochantero-femoral articulation, the parts remaining outside of
this joint are dropped before the next. moult, and are afterwards
renewed either as a straight short stump in which the articula-
y tions are already observable, or as a
miniature leg, the femur of which is
| . straight and the tibia and tarsus
Ly # i curved into a nearly complete circle ;
f 3 in the former case, the leg assumes at
Va \ the next moult the appearance that
\ \ it has in the second ease; this latter
‘ form is always changed at the succeed-
ing moult into a leg resembling the
normal limb in every respect except-
ing size, and the absence of the fourth
tarsal joint (Fig. 153). If the leg
be removed nearer to the body than
the trochantero-femoral articulation
the limb is not replaced.
The sexes are frequently ex-
tremely different; the female is usually
very much larger than the male.
This latter sex often possesses wings
when they are quite wanting in the
other sex; the resemblance to por-
tions of plants is often very much
greater in the female than it is in
the male.
We have pointed out that the
) tegmina or upper wings are usually
of small size or absent (Fig. 150,
Aschipasma catadromus), even in the
species where the lower wings are
‘ very largely developed ; in such cases
4 the latter organs are folded in a
Sita SAS, agelte reg pegs complicated, fan-like manner, and
front leg has been renewed. repose on the back, looking as if
Senegal. (After Westwood.) ’ :
they were really the tegmina (Fig.
159, Calvisia atrosignata); this appearance, moreover, is in some
species enhanced much by the fact that the part of the wing
which is outermost in the folded state is quite differently
\
ne
| ee aces | pea & oe
itl
—_
<=
——
Es
ee
i =
_ovipositor—ain the male, the modi-
| li ACRIDIIDAE wine 281
sort of embrasure filled up by a portion of the first ventral plate.
The hind body is elongate, and shows distinctly eight dorsal
segments, behind which are the
pieces forming—in the female, the
fossorial organs which replace an
fied parts connected with the ter-
minal segment. The alar organs
(Fig. 167) exhibit, according to
Brunner, the same areas as we
have described in Blattidae. Ac-
cording, however, to Redtenbacher *
the tegmina of the Acridiidae and
other saltatorial Orthoptera differ
from those of the cursorial group
(with the exception of the Phas- Fig. 167.—Alar organs of >a.
XII ACRIDIIDAE 287
the Wengern Alp with a friend,” he says, “ the grass on each side
of the path swarmed with Insects which to me rent the air with
their shrill chirruping. My friend heard nothing of this, the
Insect world lying beyond his limit of audition.” If human
ears are so different in their capacities for perceiving vibrations, it
of course becomes more probable that auditory organs so differently
constituted as are those of Insects from our own may hear sounds
when the best human ear can detect nothing audible. On the
whole, therefore, it would appear most probable that the Orthoptera
provided with acoustic organs, and which we consider dumb, are
not really so, but produce sounds we cannot hear, and do so in
some manner unknown to us. If this be the case it is probable
that these ears are special organs for hearing particular sounds.
Scudder, who has given considerable attention to the subject
of Orthopteran music, says that in N. America “the uniformity
with which each species of Stenobothrus plays its own song is
quite remarkable. One kind, Stenobothrus curtipennis, produces
about six notes per second, and continues them from one and a
half to two and a half seconds; another, S. melanopleurus, makes
from nine to twelve notes in about three seconds. In both
cases the notes follow each other uniformly, and are’ slower in the
shade than in the sun.”
Some of the species of Acridiidae, it should be noticed, produce
a noise during their flights through the air, due to the friction
of the wings; whether this has a definite importance, or whether
it may be entirely incidental, has scarcely yet been considered.
Information of a satisfactory kind as to the pogt-embryonic
development of the Acridiidae is but scanty. We have repre-
sented in Fig. 84, A, the condition in which a migratory locust,
Schistocerca peregrina, leaves the egg, and we will here complete
the account of its growth; following Brongniart,’ whose statement
is confirmed by Lestage and other naturalists. Immediately
on leaving the egg the young locust casts its skin, and is then ©
of a clear green colour, but it rapidly becomes brown, and in
twelve hours is black. At this early age the gregarious in-
stinct, possessed by this and some other species of Acridiidae,
becomes evident. In six days the individual undergoes a second
moult, after which it is black, spotted and banded with white, and
with a rose-coloured streak on each side of the hind body. The
1 Bull. Soc. Philomath. (8) v. 1893, p. 5.
288 ORTHOPTERA ‘CHAP.
third ecdysis occurs in six or eight days after the second; the
rose colour becomes more distinct, and the head is of a brown tint
instead of black. After eight days the fourth ecdysis occurs ;
the creature is then about 35 millimétres long; its colour has
much changed, the position of the markings is the same, but the
rose colour is replaced by citron yellow, the line of the spiracles
is marked with white, and at this time the creature has the “ first
rudiments of wings,” and is very voracious. In ten days another
ecdysis takes place, the yellow colour is more vivid, the prothorax
is definitely speckled with white, and the hind body is increasing
Fia. 174.—Development of wings in Caloptenus spretus: the upper row gives a lateral
view of the thoracic segments, and the lower row a dorsal view of these segments ;
1, second instar ; 2, third instar; 3, fourth instar; 4, fifth instar. (After Riley.)
zt, tegmen ; w, wing.
’
much in size. In fifteen or twenty days the sixth moult occurs,
and the Insect appears in its perfect form; the large tegmina
now present are marked with black in the manner so well known,
and the surface generally is variegated with bluish and rosy marks.
Although this is the colour in Algeria, yet apparently it is not
so farther south; the Insects that arrive thence in the French
colony are on some occasions of a different colour, viz. reddish or
yellowish, those of this latter tint being, it is believed, older
specimens of the reddish kind. M. Brongniart points out that
some Phasmidae—of the Phylliwm group—undergo an analogous
series of colour-changes in the course of the individual develop-
ment, though other species do not.
oy =. ae
f =
Mae =
ACRIDIIDAE 289
Riley and Packard have given an account * of some parts of the
post-embryonic development of the Rocky Mountain Locust, which
enables us to form a satisfactory conception of the stages of de-
velopment of the wings. Fig. 175, A, represents the first instar,
the young locust, just emerged from the egg and colourless. Fig.
174 shows some of the subsequent stages of development of the
wings, the upper line of figures giving a profile view of the
thoracic segments, and.
the lower line showing
their dorsal aspects; 1
shows the condition of
the parts in the second
instar, the chief differ-
ence from the first instar
being the development
of colour; in the third
instar there is an evident
slight development of
the future alar organs,
exhibited chiefly in the
outgrowth and lobing g&&
of the free posterior —
angles of the meso- and
metanota, as shown in
Fig. 174, 2. After the
third moult there is a
great difference; the in- .
star then disclosed —
the fourth—has under-
gone a considerable
change in the position Fie. 175.—Caloptenus spretus. North America. A,
Newly hatched, much magnified ; B, adult, natural
of the meso- and meta- ize. "(After Riley.)
thoraces, which are
thrust forward under the pronotum ; this has become more enlarged
and hood-like (Fig. 174, 3); at the same time the wing-rudiments
have become free and detached, the metathoracic pair being the
larger, and overlapping the other pair. The fifth instar (Fig. 174,
4) differs but little from the fourth, except in the larger size of the
pronotum and wing-rudiments. The sixth—shown in Fig. 175, B
1 First Ann. Rep. U.S. Ent. Comm. 1878, p. 279.
VOL. V U
290 ORTHOPTERA CHAP.
—is the perfect Insect, with the alar organs free and large, the
prothorax much changed in form, the colour different. From the
above it will be seen that the chief changes occurred at the third ~
and fifth ecdyses, after each of which a considerable difference in the
form of the Insect was revealed. In the first three instars the
sexes can scarcely be distinguished, in the fourth they are quite
distinct, and in the fifth coupling is possible, though usually it
does not occur till the final stage is attained.
The discovery that Orthoptera change their colours in the
course of their development, and even after they have become
adult, is important, not only from a physiological point of view,
but because it throws some light on the questions as to the
number of species and the geographical distribution of the
migratory locusts, as to which there has existed a great confusion.
The Acridiidae are considered to be exclusively vegetable
feeders, each individual consuming a very large quantity of food.
The mode in which the female deposits her eggs has been
described by Riley,’ and is now widely known, his figures having ©
been frequently reproduced. The female has no elongate ovi-
positor, but possesses instead some hard gonapophyses suitable for
digging purposes ; with these she excavates a hole in the ground,
and then deposits the eggs, together with a quantity of fluid, in
the hole. She prefers hard and compact soil to that which is
loose, and when the operation is completed but little trace is left
of it. The fluid deposited with the eggs hardens and forms a
protection to them, corresponding to the more definite capsules of —
the cursorial Orthoptera.
The details of the process of oviposition and of the escape of
the young from their imprisonment are of much interest. Accord-
ing to Kiinekel d’Herculais? the young Stauronotus maroccanus
escapes from the capsule by putting into action an ampulla
formed by the membrane between the head and the thorax; this
ampulla is supposed to be dilated by fluid from the body cavity,
and is maintained in the swollen condition by the Insect accumu-
lating air in the crop beneath it. In order to dislodge the lid of
the capsule, six or seven of the young ones inside combine their
efforts to push it off by means of their ampullae. The ampulla
1 Rep. Ins. Missouri, ix. 1877, p. 86.
> Bull. Soc. ent. France (6), x. 1890, p. xxxvii., and OR. Ac. Paris, cx. 1890,
p. 657.
7
i as ie ae
XII LOCUSTS 2gI
subsequently serves as a sort of reservoir, by the aid of which the
_ Insect can diminish other parts of the body, and after emergence
from the capsule, penetrate cracks in the earth so as to reach the
surface. Immediately after doing this the young Stauronotus
moults, the skin it casts being called by Kiinckel an amnios.
The cervical ampulla reappears at subsequent moults, and enables
the Insect to burst its skin and emerge from it.
The process is apparently different in Caloptenus spretus,
which, according to Riley, ruptures the egg-shell and works its
way out by the action of the spines at the apex of the tibiae.
This latter Insect when it emerges moults a pellicle, which Riley
considers to be part of the embryonic membranes. |
Riley states that a female of Caloptenus spretus makes several
egg-masses. Its period of ovipositing extends over about 62
days, the number of egg-masses being four and the total number
of eggs deposited about 100. The French naturalists have
recently observed a similar fact in Algeria, and have ascertained
that one of the migratory locusts—Schistocerca peregrina—may
make a deposit of eggs at more than one of the places it may
alight on during its migration.
It has been ascertained that the eggs of Acridiidae are very
nutritious and afford sustenance to a number of Insects, some of
which indeed appear to find in them their sole means of subsist-
ence. Beetles of the family Cantharidae frequent the localities
where the eggs are laid and deposit their eggs in the egg-masses
of the Orthoptera, which may thus be entirely devoured. Two-
winged flies of the family Bombyliidae also avail themselves of
these eggs for food, and a mite is said to be very destructive to
them in North America. Besides being thus destroyed in
enormous quantities by Insects, they.are eaten by various birds
and by some mammals,
Most of the Insects called locusts in popular language are
members of the family Acridiidae, of which there are in different
parts of the world very many species, probably 2000 being
already known. To only a few of these can the term Locust be
correctly applied. A locust is a species of grasshopper that
occasionally increases greatly in number, and that moves about. im
Swarms to seek fresh food. There are many Orthoptera that
occasionally greatly increase in numbers, and that then extend
their usual area more or less; and some
ee
?
xl | LOCUSTA OVIPOSITOR 315
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 indi-
cations 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, ec).
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 ex-
ternal rods of the ovipositor.
,
Fia. 191.—Development of ovipositor of
Locusta viridissima : a, terminal seg-
ment ; a’, cerci ; a, secondary division
of terminal segment ; 8, penultimate
(ninth) segment ; 0’, primary papillae
of this segment ; 6”, secondary divi-
sions 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.
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 0’ of the ninth segment are seen with their secondary
Fic. 192. —Structure of
ovipositor of Locusta
viridissima ; A, ar-
rangement of parts at
base, c’ being separated
and turned outwards ;
B, transverse section.
The parts of the ap-
pendage bear the same
lettering as in Fig. 191.
(After Dewitz. )
offshoots 6”; 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 LOCUSTIDAE CHAP,
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 com-
pleted structures the parts c’ have become very intimately con-
nected 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 en-
trance 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.
Fic, 193.—Ears of Locustidae: A, portion
of front leg of Odontura serricauda, 193, A) or closed by an Over-
adult ; p, prominence of integument; 7, orowth of the integument (Fig.
rim of ear; 7, tympanum ; 4, thickened ; wes
area thereof; Fu, remains of groove in 193, B), or in a condition to a
which the structure was developed. B, certain extent different from
portion of front leg of Zhamnotrizon , ‘
apterus; i, inner margin; a, slit - like either of these. The existence of
enna entre of ers i ovevaPDINE ears placed on the legs is a euri-
ous fact, but it is beyond doubt
in the Locustidae, and there is good reason for believing that analo-
gous organs exist in this situation in other Insects that have special
means of sound-production, such as the ants and the Termites.
The structure of these organs in the Locustidae has been
investigated by Graber,’ 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
1 Arch. f. mikr. Anat. xx. 1882, and xxi. See also von Adelung, Zeitschr. wiss.
Zool. liv. 1892, p. 316.
XIII EARS 7 337
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 con-
nected 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 5
not know whether the necessity ; J
for it may be connected with the T9104-Diasramof arangement of prt
respiratory system or the acoustic _ thetibiaofaLocustid. A, J,V, H, outer,
inner, anterior, posterior aspects of leg ;
organ. a,d,thin part of integument forming an-
The chief features of the acous- war rig: tee b, c, thicker portion
: of same ; 7, g, posterior tympanum ;
tic apparatus of the legs of Locus- ~ @, f and‘d, h, g, thick portions of
tidae will be gathered from the integument ; 7, %, internal protuber-
© ances of same ; /, m, n, 0, Walls of the
accompanying diagrammatic trans- _ anterior tracheal vesicle, v7; p, q 8, 7,
verse section through the tibia. In walls of the posterior tracheal vesicle,
5 y hTr ; 0”, projection of tympanal orifice
this figure the deep black parts of prothorax ; ¢r-n, tracheal inerve-
ae Oe : end organ, crista acustica ; st, rod ;
indicate the outer wall of the de, curtain-membrane ; hn, e, supra-
tibia and its prolongations, the tympanal, nerve-end organ ; in, gang-
hit aeAisata:-t t lion cells; st’, rods; ¢, point of in-
white spaces indicate 16 parts tegumental fixation of nerve endings.
filled with air, while the dotted (After Graber.)
portions are occupied by blood or some of the body organs;' the
1 The small space above Jm left free from dots is,we presume, due to an omission on the
part of Graber’s artist, but we have not thought it right to interfere with his diagram.
318 : LOCUSTIDAE | CHAP.
circular space o” is not part of the actual structure, but repre-_—
sents the area of the external acoustic orifice of the protaena
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 sur-
face, 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
XIII MUSIC 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 corre-
sponding 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 im-
portant 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 pte ts
do so, but when the wings are closed "™%,,)90-—immer face of, ase ot
? 5 tegmina of Locusta viridissima :
quite covers and conceals the musical — A, the two wing-covers separ-
; ated ; B, in natural position with
istrument. mesonotum connecting them,
The Locustidae, notwithstanding showing file and edge scraping
‘ it; a, the stridulating file; 3,
the fact that their alar organs are the rudimentary file on other
generally more ample than those of the ‘sen.
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 sustain-
ing 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 Tanana 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 Z7’h/iboscelus
camellifolius. It is very similar in appearance to Cyrtophyllus
crepitans, the Insect we have represented in Fig. 187.
The most notorious of the musical Locustids are the Katydids
320 ORTHOPTERA CHAP.
of North America. There are several species of them—they belong,
indeed, to more than one genus,—but it seems that sounds some-
what 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:' “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 rasp-
ings, 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
Fic. 196.—Katydid, Microcentrum retinerve. N. America. (After Riley.)
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.”
Pertinacity is one of the most curious features of the perform-
ance 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 per-
forms 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 chang-
1 Ann, Rep. Insects Missowri, vi. 1874, p. 159.
ere ea ee >. ,.
>
=
XIII LOCUSTIDAE 321
ing 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 some-
times with Orphania denticauda and Sarbitistes 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 variwm deposits its eggs in the galls of Cynips
in the autumn; these eggs do not hatch till the following
spring. Xiphidiwm 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 com-
pleted in the following spring, eight or nine months afterwards.
We owe to Wheeler! 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 Ken-
tucky, and in the rock-cavities of New Zealand to be one
1 Wheeler, J. Morphol. viii. 1893.
VOL, V Y
322 ORTHOPTERA CHAP,
species, although they are now referred by entomologists to
different genera.
\ AVY
| LC®
“aN je! LP
| Qe
J
g ~~
~
Fig. 197.—Dolichopoda palpata, male. Dalmatia, (After Brunner. )
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,
viz. that the tracks
and spots formed on
Fic. 198.—Leaf-like tegmen of Pterochroza ocellata: a wit
a, a, marks like those made by Insects on leaves, > leaves by the Bais. 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,
1 Verh. zool.-bot. Ges, Wien, xxxiii, 1883, p. 248.
XIII HYPERTELY 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 2 |
of a little Phaneropterid were brought
from the Soudan by the Antinori ex- a
_pedition, and have been described by
Brunner under the name of Myrmeco- |
phana fallax. The Insect is said to
bear an extraordinary resemblance to
an ant. The most peculiar feature in f
5 , an
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 Ong fee oat
_ the colour generally, the Myrmecophana “see Fallin. sti ine
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 wing-
less 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.
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 Hphippigera rugosicollis, and
tells us that the human eye can, with a little practice, acquire
the art of detecting these concealed creatures. This consists
324 ORTHOPTERA CHAP.
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 pro-
vided with means of defence of a posi-
tive nature. The Algerian Hugaster
guyont ejects two jets of a caustic
orange-coloured fluid from two pores
situate on the sides of the meso-
sternum, and covered by the anterior
coxae. This species is carnivorous as
well as herbivorous, and produces a.
sound more like humming than stridu-
lation.t |
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 Locus- —
tidae. Such a resemblance has, how-
ever, 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
Fic. 200.— Phasmodes ranatri- Insect a considerable resemblance to
avadlaans os Australia, (After the stick-Insects or apterous Phas-
midae. 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 -
1 Bonnet and Finot, Rev. Sci. Nat. (3) iv. p. 845. The word we have translated
as humming is *‘ bruissement.”
ea
XIII LOCUSTIDAE 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 repro-
duces, to a considerable extent, the form and appearance of
Proscopides or of some Tryxalides.'
\
Fic. 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 de-
1 De Saussure, Ann. Soc. ent. France, 1888, p. 151, pl. v. fig. 1.
326 ORTHOPTERA CHAP.
flexed 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’ 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.
Tia. 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 Anosto-
stoma is remarkable for its large head. Allied to it is Deinaerida
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 24 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 wing-
less, yet, as Buller informs us,’ 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
1 Indian Mus. Notes, ii. 1893, p. 172. 2 Zoologist, 1867, p. 489.
XUI LOCUSTIDAE | 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
Humegalodon and the tribe Eumegalodonidae, which is not included
in Brunner’s table of the tribes of Locustidae. The ovipositor
a
CPT ee a UPN re oie ea
1, “Ah Gia Way
Fic. 203.—Lumegalodon blanchardi, female. Borneo. x. (After Brongniart.)
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.
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 ORTHOPTERA CHAP.
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.! i
4. First two joints of the tarsi laterally smooth. (Posterior tibiae
furnished on each side with an apical spine.) Tribe -1.
PHANEROPTERIDES. (Fig. 196, Muécrocentrum; 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 prognath-
ous.) Tribe 4. Procuinipes. (Fig. 200, Phasmodes.)
5’. Foramina of the anterior tibiae forming a chink, or pro-
tected by a scale. (Fig. 193, B.)
6. Anterior tibiae with no apical spines.
7. Margins of the scrobes? 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, Copio-
phora 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.?
1 This diagnosis is an attempt to express in something apprvaching an exact
manner the distinction of the flattened from the arched or convex head.
2 Scrobes are the depressions in which the antennae are inserted.
3 There are unfortunately a few exceptions in the case of this character.
XIII LOCUSTIDAE 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. DecriciDEs.
_ 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. Elytrain the females wanting. Anterior tibiae
without apical spine on either side. Posterior tibiae
beneath with two apical spines or with none. Tribe 13.
. HeErTRopIDEs.
2’. Anterior tibiae without auditory cavities. Tegmina with no tym-
panum. Tribe 14. Gryzuacripes. (Fig. 201, Schizodactylus
monstrosus. )
1’. Tarsi distinctly compressed (most of the species apterous.) Tribe 15.
STENOPELMATIDES. (Fig. 202, Anostostoma australasiae; Fig. 197,
Dolichopoda palpata.) |
PINT a BP 8 n
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 Gryllotal-
pides). 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 possess-
. ing, 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 Gryl-
lidae 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.
Fia. 204. —House-cricket, Gryllus (Acheta) ~The family would be a
domesticus, male. ;
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
—
tO ees
CHAP. XIV f CRICKETS 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 pro-
vided 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 some-
what like the elytra of Coleoptera, and close over the-pack 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 fiat on the back
of the Insect — represents
merely the anal area, and
all the other parts must be
sought a the smaller, , de- Fic. 205.—Tegmina (sinistral) of the house-cricket.
flexed portion of the wing- A, male, inner aspect; B, female, outer
Des ’ atts aspect: a, inner margin; 0, outer margin ;
Cover. Ve saussure s Opinion, c, nervure bearing stridulating file.
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.’
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
1 See Pungur, Termes. Fiizetek, 1877, p. 223.
332 ORTHOPTERA CHAP.
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 éhe 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’ 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 produc-
1 Brunner, Verh. zool.-bot. Ges. Wien, xxiv. 1874, p. 288.
;
XIV MOLE-CRICKET 333
tion 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, Courtiliére of the French — dis placed with a few
allies in a special group, Gryllotalpides, characterised by the
lated 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,’ 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
Sseythe 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.” ;
The front legs are remark-
able structures (Fig. 206), being
beautifully adapted for burrow-
ing; 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,
Fig. 206.—Front leg of the mole-cricket.
A, outer ; B, inner aspect : @, ear-slit.
1 Natural History of Selborne, Letter xe.
334 GRYLLIDAE CHAP,
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 cowrtiliére is
supposed to be a corruption of the Latin ewrtilla. Its fondness
for the neighbourhood of water is well known. De Saussure
says that in” order to secure specimens it 1s 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, ¢).
In the allied genus Scapteriseus the tympanal membrane is, how-
ever, 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. |
The alimentary canal and digestive system of Gryllotalpa
mee
H |
:
XIV. MOLE-CRICKET 335
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, ¢); 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 Mal-
pighian tubules open is seated
near the commencement of
the small intestine. These
tubules are very fine, and are
about one hundred in num-
ber. The arrangement by
which the Malpighian tub-
ules open into a common duct
instead of into the intestine
itself appears to be charac-
teristic of the Gryllidae, but
is said to occur also in
Ephippigera, a genus of
Locustidae. According to
Leydig! and Schindler the
Malpighian tubules are of
two kinds, differing in colour,
and, according to Leydig, in
contents and histological Fic. 207.—Alimentary canal and appendages of
; the mole-cricket: a, head; 0, salivary
structure. Near the posterior glands and receptacle ; c, lateral pouch ; d,
extremity of the rectum stomsto-gastric nerves ; ¢, ea lobes of
stomach ; f, peculiar organ; g, neck of
there is a lobulated gland stomach ; h, plicate portion of same; 7%, rec-
having a reservoir connected = "i be traps sg . pieces! decal ;
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 ;
1 Miiller’s Arch. 1859, p. 159.
yy
336 ORTHOPTERA "CHAP.
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,’ 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? and Korotneff? 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 con-
dition of the digestive organs that
necessitates the unusual care the
mother bestows on her young.
The genus Cylindrodes (Fig. 208,
C. kochi) comprises some curious and
rare Insects of elongate, slender form.
They are natives of Australia, where
i the first species known of the genus
Fic. 208. — Cylindrodes kocht. was found in Melville Island by Major
Australia. A, outline of the
Insect with five of the legs and Campbell, from whom we learn that
the extremity of the body muti- these Insects burrow in the stems of
lated ; B, middle leg. (After ‘
de Saussure.) plants, and are so destructive that he
was unable to keep a single plant in
his greenhouse on account of the ravages of Cylindrodes
campbell. The form of these Insects is beautifully adapted to
1 Bull. Soe. ent. France, 1893, p. ceexli.
2 Zeitschr. wiss. Zool. xxiii. 1876, p. 122. 3 Ibid, xli. 1885, p. 570.
.
XIV TRIDACTYLIDES 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 7ridactylus is considered by de Saussure to form,
with its ally Rhipipteryx, a division of Gryllotalpinae, but ney
are treated, perhaps more
correctly, by Brunner as a
separate tribe. TZ. varie-
gatus (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 an- /f
tennae. Its alar organs
are imperfect, and not like
Fic, 209.—Tridactylus variegatus, France.
those of other Gryllidae in either form or neuration. The hind
legs are of peculiar structure, the tibiae terminating in two pro-
cesses 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
ficure. 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’ 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.
The exotic genus Rhipipteryx is allied to Tridactylus. It is
widely distributed in South America, but the little Insects that
1 Morph. Jahrb. xv. 1889, p. 400.
VOL. V Z
338 ORTHOPTERA CHAP,
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 Ama-
zonian species we figure (Fig. 210), the
wings, instead of being mere rudiments,
as in 7'ridactylus, 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.
Fig. 210.—Rhipipteryz sp., The Tridactylides have no tympana on
AIBN OEE the legs, and their affinity with the Gryl-
lidae is very doubtful. Dufour thought 7. variegatus to be more
allied to the Acridiidae. He based this opinion chiefly on some
points of the internal anatomy, but pointed out that Zridactylus
differs from the Acridiidae in having no air-sacs in the
body.
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 pre-
dilection 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 bake-
a. de
Locustidae (Fig. 189) and Mantidae
a _ GRYLLIDAE 339
houses. Usually the wings in the cricket are elongate, and pro-
ject 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 Mediter-
ranean region, the male of which has
the head prolonged into a curious pro-
cess (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 extra-
ordinary structures we have figured in ¢
A
(Fig. 136), though it appears impossible
. Fia. 211.—Platyblemmus lusi-
to treat the cephalic appendages of Platy- sities wenlee We Gent or
blemmus as ornamental objects; their head; B, profile of Insect
‘ ; : with most of the appendages
import is at present quite obscure. valiovad.
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 ex-
panded. 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
mnany specimens. Some species are always micropterous.
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 GRYLLIDAE CHAP. XIV
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, 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, Rhipipterysx 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. MyrmeEco-
PHILIDES.
4’. Posterior tibiae biseriately spinose. Ovipositor straight.
5, Antennae short, thickish, almost thread-like. Facial
scutellum exserted between antennae. Posterior tibiae
dilated. Gen. Myrmecophila,?
5’. Antennae elongate, setaceous. Facial scutellum trans-
verse, 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, Platy-
blemmus lusitanicus.)
6’. Posterior tibiae slender, armed with slender spines,
and serrate between them. Tribe. 5. OxcAN-
THIDES.
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.
* Mem. Soc. phys. Genéve, xxv. 1877, and Biol. Centr. Amer. Orthoptera, 1894, p. 198.
* The genus Myrmecophila, being exceptional in several respects, is treated separately.
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 com-
plez minute mesh-work. (One division, Mallophaga, con-
sists entirely of wingless forms ; in TLermitidae 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.
ee Y
AS Ms
s sa> >! eee! cans
FEIN an N
ee: are
es, E95 d
LITE 5
Kg
Fic. 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 NEUROPTERA CHAP.
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.
. 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.
. Trichoptera.—Development as in division 4. Mandibles absent in the adult
Insect. Life aquatic in the early stages. Fam. 11. Phryganeidae.
ww
on
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 Phas-
midae 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 Pseudo-
neuroptera; this sub-Order comprised our Termitidae, Psocidae,
Eplemeridae, 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
XV FOSSIL NEUROPTERA 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. Brong-
niart’s very important work! on the Insects of that epoch has
appeared. He considers that these ancient fossils may be classi-
fied 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 Orthop-
tera, 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 inter-
pret as the actual ancestors, of our existing Ephemeridue, Odonata,
and Perlidae.
Some of the fossils restored and described by the French ento-
mologist are of great interest. We shall notice the Prote-
phemerides, 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,
1 Insectes fossiles des temps primatires, 1893, vol. i. and atlas.
344 NEUROPTERA CHAP. |
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 consider-
ably 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, Cory-
daloides scudderi, is of great interest, as it is believed by Brong-
niart 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. 215); these are looked
on as analogous to the ex-
pansions of meso- and meta-
thorax, which are supposed by
some writers to have been
Fic, 213.—Lithomantis carbonaria. Car- the rudiments from which
boniferous strata of Commentry, France. wings were developed. These
Anes epneniaes 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.
The third family of extinct carboniferous Neuroptera is the
Stenodictyopterides, in which Brongniart places the Dictyoneura of
> tee
“Nat
—-
ime Nh Re Peo: J nent time,
XV MALLOPHAGA 345
*
Goldenberg, the North American Haplophlebiwm, 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 car-
boniferous Neuroptera have now been discovered, no larvae or
immature forms have been found.
We now pass to the consideration of the divisions of Neurop-
tera still living. }
Fam. I. Mallophaga—Bird-Lice or Biting Lice.
Small Insects, wingless, with large head; thorax usually cf two,
rarely of one or three segments ;
prothorax always distinct ; hind
body consisting of eight to ten
segments, in addition to the pos-
terior two thoracic segments which
usually are but little or not at
all separated from it. The meta-
morphosis 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 ; Fic. 214.—Trinoton luridum.
they rarely come quite to the surface, Lives on the common duck
so)-that they are not detected on a 4 Various species of Anas.
(After Giebel.)
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.
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
346 _ MALLOPHAGA CHAP.
under side of the head. 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 situ-
ated 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 scrap-
ing instrument (o/, Fig.
Fic. 215.—Under-surface of head of Lipeurus 215). The mandibles are
heterographus. (After Grosse.) ol, Labrum ; sharply toothed and appar-
md, mandible ; mx, maxilla; ul, labium. i ;
| ently act as cutting instru-
ments. The maxillae have been described in the principal
work on the family’ as possessing in some cases well-developed
palpi. According to Grosse? 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 undi-
vided; on each side
We . Fic. 216.—Under lip of Nirmus, A; and of Tetroph-
there ne paraglossa thalmus chilensis, B. (After Grosse.) m, Mentum ;
bearing an oval pro- g, ligula ; pl, palp ; pg, paraglossa ; hy, lingua.
cess, 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,
1 Giebel and Nitzsch, Jnsecta epizoica, folio, 1874.
2 Zeitschr. wiss. Zool. xiii. 1885, p. 537.
weXV | MALLOPHAGA 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 Zvrinoton (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 repro-
ductive 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 Sf Yry a
any species. & Res ~
The nervous system has been Ned ;
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 evi-
dently of two conjoined halves. The
alimentary canal has a slender, elon- ye, 217,.—Ganglia of nervous sys-
gate oesophagus, dilated behind into a ite, eee enn i aad
crop; this is frequently received be-
tween two cornua formed by the anterior part of the stomach,
_ which, except for these, is simply tubular in form, though some-
what 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 MALLOPHAGA CHAP,
There are only four Mal-
pighian 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 exist-
ing 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 pro-
ceeding from the bicapsular vesicula semi-
posterior portion of the oesophagus...
Fic. 218.—Alimentary canal
of Docophorus fuscicollis.
(After Giebel.) a, Oeso-
phagus ; 6, paunch; a’,
posterior division of oeso-
phagus ; ¢, chylific ven-
tricle or stomach ; d,'Mal-
pighian tubes; e, small
intestine ; jf, glandular
girdle ; g, rectum.
nalis; these structures have been figured
by Grosse’ 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 recepta-
culum seminis.
The eggs of some Mallophaga have been figured by Melnikow ;?
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 ex-
tremely like the adult, and are thought to moult frequently ; ths
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 Liseha: 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
1 Zeitschr. wiss. Zool. xiii. 1885, pl. xviii. f. 15.
2 Arch. f. Naturg. xxxv. i. 1869, p. 154, pls. x. xi.
Le
EE
xv. MALLOPHAGA 349
ina 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.
Fic. 219.—Lipeurus ternatus, male ; Fia. 220.—Trichodectes latus, male ;
inhabits Sarcorhamphus papa. inhabits the dog, Canis famili-
(After Giebel.) aris.
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, Doco-
phorus 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 MALLOPHAGA cua.
221), which is said to have been figured by Redi two hun-
dred years ago under the name of Pulex
capt. This species multiplies to a con-
siderable 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 per-
Be haps have been really due to itch-mites.
inhabits the common fowl, Lhere is, however, no doubt that this
sa domesticus, (After snecies may infest poultry, especially if
Re 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.
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.t 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 (Z'richodectes, Fig. 220; and Gyropus)
belong to the two chief divisions of Mallophaga. The genus
Menopim 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.”
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
1 Op. cit. pp. vii.-xiv. For classification, ete., see also Piaget, Les Pédicudlines.
Leyden, 1880. * Zeitschr. wiss. Zool. xiii. 1885, p. 532.
xv EMBIIDAE 351
by means of the parasitic two-winged flies that infest birds.
The writer has recorded! 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.
Hlongate feeble Insects ; with small prothoraz, elongate meso- and
meta - thorax, which
may either bear wings
or be without them.
In the former case
these organs are not
caducous, are deli-
cately membranous,
and all of one consist-
ence, with three or
four indefinite longi-
tudinal nervures and
-veinle Fic. 222.—Oligotoma michaeli. (After
a few cross-veinlets. ds)
hy, | y
MEL Di a
AS ae Up
rt
. i }
. m» A
fj
we 17S
i
4 di
The development 1s
incompletely known. The individuals do not form organised
societies.
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
1 P, ent. Soc. London, 1890, p. xxx.
352 NEUROPTERA CHAP.
it is not a matter for surprise that little should be known about
them.
The simple antennae are formed of numerous joints, probably
varying in number from about fifteen to twenty-four. The mouth
is mandibulate. Chatin states’ that the pieces homologous with
those of a maxilla can be detected in the
mandible of Hmbia. 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 abdo-
men 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 develop-
ment of the meso- and meta-thorax and
their elongation in comparison to the pro-
thorax. When they bear wings there is no
modification or combination of the segments
Fic, 223.—Under-surface for the purposes of flight, the condition of
of Embia sp. Andalusia. : ; ee
these parts being, even then, that of wing-
less Insects; so that the Embiidae that have wings may be
described as apterous-
like Insects provided
with two pairs of in-
efficient 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 wing ; B, outline of the wing, showing nervures.
appendage between (After Wood-Mason.) 1, Costal; 2, subcostal; 3,
radial ; 4, discoidal ; 5, anal nervure.
the claws. Ai Sian"
The wings in Embiidae are very peculiar; they are extremely
1 Bull. Soc. Philom. (7) ix. p. 33.
Fig. 224.—Anterior wing of Oligotoma saundersii: A, the
ee
f * + 4
=!
ted Wess
wes?
XV EMBIIDAE 353
flimsy, and the nervures are ill-developed; stripes of a darker
brownish colour alternate with pallid spaces. We figure the an-
terior wing of Oligotoma saundersit, 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 inter-
pretation of the nervures. He considers! that the brown bands
“mark the original courses of veins which have long since dis-
appeared.” A similar view is taken by Redtenbacher,? 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* there are ten pairs of stigmata, two
thoracic and eight abdominal; these are connected by longi-
tudinal and transverse tracheae into a single system. The
ganglia of the ventral chain are, one suboesophageal, three thor-
acic, 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 Japya#, there being in
each five short egg-tubes, opening at equal intervals into a
straight duct. The testes are remarkably large; each one con-
sists 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
1 P. Zool. Soc. London, 1883, p. 628.
2 Ann. Hofmus. Wien, i. 1886, p. 171.
3 Atti Acc. Gioenia, vii. 1893.
VOL, V 2A
354 NEUROPTERA CHAP,
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’ as possess-
ing 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 uncer-
tain. These Insects have been detected in various parts of the
world. In the Sandwich Islands Oligotoma insularis was dis-
covered by the Rey. T: Blackburn in the wood and thatch form-
ing the roofs of natives’ houses. A species has been found in
Prussian amber, and Grassi thinks that Hmbia_ solieri—one of
the Mediterranean species—is not to be distinguished with cer-
tainty 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
1 J. Linn. Soe, Zool. xiii. 1878, pl. xxi. f. 2.
XV ! EMBIIDAE 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 assist-
ance 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.
Considerable difference of opinion has prevailed as to the
allies of thes? 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 ordi-
nary Orthoptera. Brauer places the family in his Orthoptera
genuina.
1 Canadian Entomologist, xvii. 1885, throughout.
CHAPTER XVI
NEUROPTERA CONTINUED—TERMITIDAE, TERMITES OR WHITE ANTS
Fam. III. Termitidae—White Ants, Termites.
Each species is social, and consists of winged and wingless indi-
viduals. 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
_
Fic, 225.—Termes (Hodotermes) mossambicus. Winged adult. (After Hagen.)
extend far beyond the apex of the abdomen; the hind pair
‘as remarkably similar in size, form, and consistence to the
JSront 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 re-
posing 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
CHAP. xXVl TERMITIDAE 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 mem- .
branous 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 accom-
panied by facetted eyes. The antennae »,, 926.— Termes bellicosus.
are simple, consisting of from nine to Labium, A, maxilla, B, of
° fs : ; winged adult ; lower face of
thirty-one joints, which differ but gach. (After Hagen.)
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,
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 pro-
358 NEUROPTERA CHAP.
notum, 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 accord-
ing 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 volumi-
nous, 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. ;
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 ob-
servers of the Insects; these con-
vulsive movements may possibly
Fic. 227.—Front tibia and tarsus of be connected a ahh thie produpiion
Calotermes rugosus larva, showing of sound, which may perhaps be
Millen} organ, x 90. (After F. evoked by contact between the
back of the head and the pro-
notum ; 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 Miiller, and we
reproduce (Fig. 227) one of his figures. The structure seems to
1 Jena. Zeitschr. Naturw. ix. 1875, pl. xii. See also Stokes in Science, xxii.
1893, p. 273.
XVI TERMITIDAE 359
be in plan and position similar to the ear of Locustidae,
though much less perfect.
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 differ-
ences in the extent to
which they are made up
of the various fields. This ,
is shown in Fig. 228, 2%
where the homologous * >
nervures are numbered
according to-the systems
of both Hagen and Red-
tenbacher. The area, VII,
that forms the larger part
; ; »o_. FIG. 228.—Wings of Termites: A, Termes lucifugus ;
of the Beeee Ul C, ore B, Hodotermes brunneicornis; C, Culotermes
spondstothesmall portion — nodulosus. (After Redtenbacher: B and C
: diagrammatic.) III, V, VII, homologous areas
at the base of the mane and nervures according to Redtenbacher. 1,
m4 B; The most re- Costal ; 2, subcostal ; 3, median; 4, submedian
markable feature of the nervures according to Hagen,
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 detach-
ment 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’ 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 coales-
cence 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 Termi-
tidae 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
1 Ann. Hofmus. Wien, i. 1886, p. 183.
360 NEUROPTERA , CHAP.
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. Miiller." The number of spiracles is ten; Hagen says three
thoracic and seven abdominal, Miiller- 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. . Salivaky
glands exist, as also do salivary reservoirs; these latte 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 import-
ance in the economy of Termites.
‘These creatures emit minute quanti-
Fic. 229.—Head and Savane ties of a secretion that is corrosive, and
canal of Termes lueifugus can act on metal and even glass; its
(nymph). a, head ; 4, salivary
glands ; ¢, salivary receptacles; Nature and source are not understood.
d, crop; @ stomach ; Jf, intes- Hagen describes peculiar structures in
tinal paunch; g, small, h, ; ac mika 4
large intestine ; 7, Malpighian the rectum to which he is inclined ®
» tubes ; &, extremity of body. to ascribe the origin of this substance,
(After Dufour.) he ;
but this is very uncertain,
The brain is small; the infra-oesophageal ganglion is placed
ty
1 Jena. Zeitschr, Naturw. ix. 1875, p. 257.
2 Bidie, in Nature, xxvi. 1882, p. 549, * Linnaea Entomologica, xii. 1858, p. 305.
XVI ; TERMITIDAE 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 con-
tinuance 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 com-
munity 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 NEUROPTERA CHAP.
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? of a swarm of 7. 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 dispropor-
tionately large head, and are of a peculiar white colour as if
filled with milk. This condition of milkiness they retain, not-
withstanding 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 develop-
ment; 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 under-
stand 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 Ter-
mites 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.
Calotermes flavicollis inhabits the neighbourhood of the
Mediterranean Sea; it is a representative of a large series of
species in which the peculiarities of Termite life are exhibited
1 P. Boston Soc. xx. 1878, p. 118.
i
:
a TERMITIDAE 363
in a comparatively simple manner. There is no special caste of
workers, consequently such work as is done is carried on by the
other members of the community, viz. soldiers, and the young
and adolescent. The habits of this species have recently been
studied in detail in Sicily by Grassi and Sandias! The Insects
dwell in the branches and stems of decaying or even dead trees,
where they nourish themselves on those parts of the wood in
which the process of decay is not far advanced; they live in the
interior of the stems, so that frequently no sign-of them can be
seen outside, even though they may
be heard at-work by applying the
ear to a branch. They form .no
special habitation, the interior of
the branch being sufficient protec-
tion, but. they excavate or increase
the natural cavities to. suit their
purposes. It is said that they
line the galleries with proctodaeal
cement; this is doubtful, but they
form barricades and _ partitions
where necessary, by cementing
together the proctodaeal products py, 939,—gome individuals of Calo-
with matter from the salivary _— termes flavicollis: A, nymph with
glands or regurgitated from the aie rial vino Mi
anterior parts of the alimentary dividual. (After Grassi.)
canal. The numbers of a com- :
munity only increase slowly and remain always small; rarely do
they reach 1000, and usually remain very much below this. The
king and queen move about, and their family increases but slowly.
After fifteen months of their union they may be surrounded by
fifteen or twenty young; in another twelye months the number
may have increased to fifty, and by the time it has reached some
five hundred or upwards the increase ceases. This is due to the
fact that the fertility of the queen is at first progressive, but
ceases to be so. A queen three or four years old produces at
the time of maximum production four to six eggs a day. When
the community is small—during its first two years—the winged
individuals that depart from it are about eight or ten annually,
but the numbers of the swarm augment with the increase of the
1 4tti Acc. Gioen. vi. and vii. 1893 and 1894.
364 ) NEUROPTERA CHAP,
population. The growth of the individuals is slow; it appears —
that more than a year elapses between the hatching of the egg
and the development of the winged Insect. The soldier may
complete its development in less than a year; the duration of its
life is not known; that of the kings and queens must be four or
five years, probably more. After the winged Insects leave the
colony they associate themselves in pairs, each of which should,
if all goes well, start a new colony.
The economy of Termes lucifugus, the only European Termite
besides Calotermes flavicollis, has been studied by several
observers, the most important of whom are Lespés’ and Grassi
and Sandias. This species is much more advanced in social life
than Calotermes is, and possesses both workers and soldiers —
(Fig. 231, 2, 3); the individuals are much smaller than those of —
Calotermes. Burrows are made in wood of various kinds, furni-
ture being sometimes attacked. Besides making excavations
this species builds galleries, so that it can move from one object
to another without being exposed; it being a rule—subject to
certain exceptions—that Termites will not expose themselves
in, the outer air. This is probably due not only to the
necessity for protection against enemies, but also to the fact
that they cannot bear a dry atmosphere; if exposed to a drying
air they speedily succumb. Occasionally specimens may be seen
at large; Grassi considers these to be merely explorers. Owing
to the extent of the colonies it is difficult to estimate with
accuracy the number of individuals composing a community, but
it 1s doubtless a great many thousands. Grassi finds the economy
of this species in Sicily to be different from anything that has
been recorded as occurring in other species; there is never a true
royal pair. He says that during a period of six years he has
examined thousands of nests without ever finding such a pair.
In place thereof there are a considerable number of complementary
queens—that is, females that have not gone through the full
development to perfect Insects, but have been arrested in various
stages of development. In Fig. 231, Nos. 4 and 5 show two of
these abnormal royalties; No. 4 is comparatively juvenile in
form, while No. 5 is an individual that has been substituted in
an orphaned nest, and is nearer to the natural condition of perfect
development. We have no information as to whether any develop-
1 Ann. Set. Nat. Zool. (4) v. 1856, p. 227. .
XVI TERMITIDAE 3 365
ment goes on in these individuals after the state of royalty is
assumed, or whether the differences between these neoteinic queens
are due to the state of development they may happen to be in
when adopted as royalties. Kings are not usually present in
these Sicilian nests; twice only has Grassi found a king, but
Fig. 231.—Some of the forms of Termes lucifugus. 1, Young larva; 2, adult worker ;
3, soldier ; 4, young complementary queen; 5, older substitution queen ; 6, per-
fect winged Insect. (After Grassi.)
he thinks that had he been able to search in the months of
August’ and September he would then have found kings. It
would appear therefore that the complementary kings die, or are
killed after they have fertilised the females. Parthenogenesis
is not thought to occur, as Grassi has found the spermathecae of
the complementary queens to contain spermatozoa.
366 NEUROPTERA CHAP. XVI~
The period of development apparently occupies from eighteen
to twenty-three months. At intervals swarms of a great number
of winged individuals leave the nest, and are usually promptly
eaten up by various animals. After swarming, the wings are
thrown off, and sometimes two specimens or three may be seen
running off together; this has been supposed to be preliminary
to pairing, but Grassi says this is not the case, but that the
object is to obtain their favourite food, as we shall mention
subsequently. | :
Although these are the usual habits of Zermes lucifugus at
present in Sicily, it must not be concluded that they are invari-
able; we have in fact evidence to the contrary. Grassi has
himself been able to procure in confinement a colony—or rather
the commencement of one—accompanied by a true royal pair;
while Perris has recorded’ that in the Landes he frequently -
found a royal pair of TZ. lucifugus under chips; they were
accompanied in nearly every case by a few eggs. And Professor ~
Perez has recently placed a winged pair of this species in a box
with some wood, with the result that after some months a young
colony has been founded. It appears probable therefore that
this species at times establishes new colonies by means of royal
pairs derived from winged individuals, but after their establish-
ment maintains such colonies as long as possible by means of
complementary queens. It is far from improbable that distinc-
tions as to the use of true and complementary royalties may be
to some extent due to climatic conditions. In some localities
T. lucifugus has multiplied to such an extent as to be very
injurious, while in others where it is found it has never been
known to do so.
The Termitidae of Africa are the most remarkable that have
yet been discovered, and it is probably on that continent that the
results of the Termitid economy have reached their climax. Our
knowledge of the Termites of tropical Africa is chiefly due to
Smeathman, who has described the habits of several species,
among them 7’ bellicosus. It is more than a century since
Smeathman travelled in Africa and read an account of the
Termites to the Royal Society. His,information was the first
of any importance. about Termitidae that was given to the
world; it is, as may be well understood, deficient in many
1 Ann. Soc. ent. France (5), vi. 1876, p. 201. 7 Phil. Trans. xxi. 1781, pp. 139-192.
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368 NEUROPTERA "CaP.
details, but is nevertheless of great value. Though his state-
ments have been doubted they are truthful, and have been
confirmed by Savage. 7. bellicosus forms buildings compar-
able to human dwellings; some of them being twenty feet in
height and of great solidity. In some parts of West Africa
these nests were, in Smeathman’s time, so numerous that they
had the appearance of villages. Each nest was the centre of
a community of countless numbers of individuals; subter-
ranean passages extended from them in various directions. The
variety of forms in one of these communities has not been well
ascertained, but it would seem that the division of labour is
carried to a great extent. The soldiers are fifteen times the size
of the workers. The community is dependent on one royal
couple. It is the opinion of the natives that if that couple perish
so also does the community; and if this be correct we may
conclude that this species has not a perfect system of replacing
royal couples. The queen attains an almost incredible size
and fertility. Smeathman noticed the great and gradual growth
of the abdomen, and says it enlarges “to such an enormous size
that an old queen will have it increased so as to be fifteen
hundred or two thousand times the bulk of the rest of her body,
and twenty or thirty thousand times the bulk of a labourer, as
I have found by carefully weighing and computing the different
states.” He also describes the rate at which the eggs are pro-
duced, saying that there is a constant peristaltic movement” of
the abdomen, “so that one part or other alternately is rising
and sinking* in. perpetual succession, and the matrix seems never
at rest, but is always protruding eges to the amount (as I
have frequently counted in old queens) of sixty in a minute, or
eighty thousand and upward in one day of twenty-four hours.”
_ This observer, after giving an account of the great swarms of
perfect winged Insects that are produced by this species, and after
describing the avidity with which they are devoured by the
Hymenopterous ants and other creatures, adds: “ I have discoursed
with several gentlemen upon the taste of the white ants; and on
comparing notes we have always agreed that they are most
1. Ann. Nat. Hist. (2) v. 1850, p. 92.
2’Dr. G. D. Haviland informs the writer that he thinks it probable this so-called
peristaltic movement is merely the result of alarm ; he has not, however, had any
apportunity of observing 7’. belZicosus.
.
XVI. TERMITIDAE 369
delicious and delicate eating. One gentleman compared them to
sugared marrow, another to sugared cream and a paste of sweet
almonds.”
From the preceding brief sketch of some Termitidas we may
gather the chief points of importance in which they differ from
other Insects, viz. (1) the existence in the community of in-
dividuals—workers and soldiers—which do not resemble their
parents; (2) the limitation of the reproductive power to a single
pair, or to a small number of individuals in each community, and
the prolongation of the terminal period of life. There are other
social Insects besides Termitidae: indeed, the majority of social
Insects—ants, bees, and wasps—belong to the Order Hymen-
optera, and it is interesting to note that analogous phenomena
occur inthem, but nevertheless with such great differences that
the social life of Termites must. be considered as totally distinct
from that of the true ants and other social Hymenoptera.
Development.—Social Insects are very different to others not
only in the fact of their living in society, but in respect of
peculiarities in the mode of reproduction, and in the variety of
habits displayed by members of a community. The greatest
confusion has arisen in reference to Termitidae in consequence
of the phenomena of their lives having been assumed to be
similar to those of Hymenoptera; but the two cases are very
different, Hymenoptera passing the early parts of their lives as
helpless maggots, and then undergoing a sudden metamorphosis to
a totally changed condition of structure, intelligence, and instinct.
The development of what we may look on as the normal form
of Termitidae—that is, the winged Insects male and female—is
on the whole similar to that we have sketched in Orthoptera ; the
development in earwigs being perhaps the most similar. The
individuals of Termitidae are, however, in the majority of cases
if not in all, born without eyes; the wing-rudiments develop
from the thoracic terga as shorter or longer lobes according to
_ the degree of maturity ; ; as in the earwigs the number of joints
in the antennae increases as development advances. All the young
are, when hatched, alike, the differences of caste appearing in the
course of the subsequent development; the most important of
these differences are those that result in the production of two
special classes—only met with in social Insects—viz. worker
and soldier. Of these the workers are individuals whose develop-
VOL, V 28
370 NEUROPTERA CHAP.
ment is arrested, the sexual organs not going on to their full
development, while other organs, such as the eyes, also remain
undeveloped ; the alimentary canal and its adjuncts occupy nearly
the whole of the abdominal cavity. The adult worker greatly
resembles—except in size—the young. Grassi considers that the
worker is not a case of simple arrest of Sa Mages but - that
some deviation accompanies the arrest.
The soldier also suffers an arrest of development in certain
respects similar to the worker; but the soldier differs'in the im-
portant fact that the arrest of the development of certain parts
is correlative with an extraordinary development of the head,
which ultimately differs greatly from those of either the ‘worker
or of the sexual males and females,
Soldier.—All the parts of the head of the oldie undergo a
greater or less change of form; even
the pieces at its base, which connect
it by means of the cervical sclerites
with the prothorax, are altered. The
parts that undergo the greatest modi-
fication are the mandibles (Fig. 233,
B); these become much enlarged in size
and so much changed in form that in
a great many species no resemblance.
to the original shape of these organs
can be traced. It is a curious fact that.
the specific characters are betterexpressed
in these superinduced modifications than
E they are in any other part of the
organisation (except, perhaps, the wings).
The soldiers are not alike in any two
‘species of Termitidae so far as “we
D
know, and it séems impossible to ascribe
the differences that exist between. the
Fic. 238.—The pairs of mandi- soldiers of different species of Termitidae
bles of different adult indi- to special adaption for the work they
id: " - t . Me
Suigepore, *. OF peti, have to perform. Such a suggestion 1s
B, bas “seat oF. dee justifiable only in the case of the: Nasuti
Ne NINE AINE’ (Fig. 23.4, 1), where the front of the head
is prolonged into a point: a duct opens at the extremity of
this point, from which is exuded a fluid that serves as a cement for
‘SOLDIERS. 37a
- constructing the nest, and is perhaps also used to disable enemies.
_ Hence the prolongation and form of the head of these Nasuti may
be fairly described as adaptation to useful ends. As regards the —
_ great variety exhibited by other soldiers—and their variety is
- much greater than it is in the Nasuti—it seems at present im-
_ possible to treat it as being cases of special adaptations for useful
_ purposes. On the’ whole it would be more correct to say
CU
\
Fic. 234.—Soldiers of different species of Termites. (After Hagen.) 1, Termes armiger ;
2, T. dirus; 3, Calotermes flavicollis; 4, T. bellicosus; 5, T. occidentis; 6, T.
cingulatus (2) ; 7, Hodotermes quadricollis (?); 8, T. debilis(?), Brazil.
j that the soldiers are very dissimilar in spite of their having to
_ perform similar work, than to state that they are dissimilar in
conformity with the different tasks they carry on.
The Termite soldier is a phenomenon to which it is difficult
_ to find a parallel among Insects. The soldier in the true ants
is usually not definitely distinguished from the worker, but it is
possible that in the leaf-cutting ants, the so-called soldier may
_ prove to be more similar in its nature to the Termite soldier,
_ The soldiers of any one species of Termite are apparently ex-
:
372: NEUROPTERA CHAP,
tremely similar to one another, and there are no intermediates
between them and the other forms, except in the stages of
differentiation, But we must recollect that but little is yet
known of the full history of any Termite community, and it is
possible that soldiers which in the stage of differentiation promise
to be unsatisfactory may be killed and eaten,—indeed there is some
evidence to this effect. . There is too in certain cases some difference
—larger or smaller size being the most important—between the
soldiers of one species, which may possibly be due to the different -
stage of development at which their differentiation commenced.
It would at present appear that, notwithstanding the remark-
able difference in structure of the soldiers and workers of the
white ants, there is not a corresponding difference of instinct.
It is true that soldiers do more of certain things than workers
do, and less of others, but this appears to be due solely to their
possession of such very different structures; and we are repeatedly
informed by Grassi that all the individuals in a community take
part, so far as they are able, in any work that is going on, and
we find also in the works of other writers accounts of soldiers
performing acts that one would not have expected from them.
The soldiers are not such effective combatants as the workers are.
Dudley and Beaumont indeed describe the soldiers as merely look-
ing on while the workers fight So that we are entitled to con-
clude that the actions of the soldiers, in so far as they differ from
those of the rest of the community, do so because of the different —
organisation and structures of these individuals. We shall, when
speaking of food, point out that the condition of the soldier in
relation to food and hunger is probably different from that of
the other forms.’ 3 ; Of
Various Forms of a Community.—tThe soldiers and workers
are not the only anomalous forms found in Termitid communi-
ties; indeed on examining a large nest a variety of forms may
be found that is almost bewildering. Tables have been drawn
up by Grassi and others showing that as many as fifteen kinds
may be found, and most of them may under certain circumstances
coexist. Such tables do not represent the results of actual
examination in any one case, and they by no means ade-
quately represent the number that, according to the most recent
observations of Grassi, may be present; but we give one taken
1 Tr, N. York Ac, viii. 1889, pp. 85-114; and ix. 1890, pp. 157-180.
i
F
les)
Q
XvI¥ TERMITIDAE 373
from Grassi, as it conveys some idea of the numerous forms that
exist in certain communities. .In this table the arrangement,
according to A, B, C, D, E, represents successive stages of the
Beretopment + — :
Forms of Reeves. fosfuguss (After Grassi.) Zool. Anz. xii. 1889, p. 360.
1. Young, undifferentiated larvae.
| | |
g | |
2. Larvae that will 3. Larvae that will 4, Reserves for royal specimens:
not mature the sexual mature the sexual (only present when 14, 15, and 11
organs, - organs. are wanting, or when 14and ld are
present in insufficient numbers).
. |.
| |
5. Larvae of 6. Pie il of - 9, Nymphs of the - 10. Nymphs of - LT. Reserves for
soldiers, workers. first form. the second form, royal pairs (only
| present when 14,
15, and 4 are want-
ing, or when the
two latter are
present in insuffi-
7. Soldiers. '8. Workers. cient numbers).
Wie 12. Winged 13. Reserve
Insects. royal pairs ?
14: poral 16. Substi tution
couple. . " royal pairs.
On inspecting this table it will be perceived that the variety of
forms is due to three circumstances—(1) the existence of castes
that are not present in ordinary Insects; (2) the coexistence of
young, of adolescents, and of adults; and (3) the habit the Termites
have of tampering with forms in their intermediate stages, the
result of which may be the substitution of neoteinic individuals in
place of winged forms.
This latter procedure is far from being eamen lace understood,
but to it are probably due the various abnormal forms, suchas
soldiers with rudiments of wings, that have from timé to time
been discovered in Termite. communities, and have given rise to
-much perplexity.
- ~ In connexion with this subject we may call attention to the
necessity, when examining Termite nests, of taking cognisance
of the fact that more than one species may be present. Bates
found different Termites living together in the Amazons Valley,
and Mr. Haviland has found as many as five species of Termitidae
and three of true ants in a single mound in South Africa. In
this latter case observation showed’ that, though in -such close
proximity, there was but little further intimacy between the
‘species. There are, however, true inquiline, or guest, Termites,
374 NEUROPTERA CHAP.
of the genus Hutermes, found in various parts of the world living
in the nests of other Termitidae.
Origin of the Forms.—The interest attaching to the various
forms that exist in Termites, more particularly to the worker and
soldier, is evident when we recollect that these never, so far as
we know,. produce young. In the social Hymenoptera it has
been ascertained that the so-called neuters (which in these
Insects are always females) can, and occasionally~do, produce
young, but in the case of the Termites it has never been sug-
gested that the sexual organs of the workers and soldiers, whether
male or female, ever become fruitful; moreover, the phenomena
of the production of young by the white ants are of such a nature
as to render it in the highest degree improbable that either —
workers or soldiers ever take any direct part in it. Now the
soldier is extremely different from the sexual individuals that
produce the young, and seeing that: its peculiarities are not, in
the ordinary sense of the word, hereditary, it must be of great
interest to ascertain how they arise. .
Before stating the little information we possess on this sub-
ject, it is necessary to reiterate what we have already said to the
effect that the soldiers and workers are no% special to either sex,
and that all the young are born alike. It would be very natural
to interpret the phenomena by supposing the workers to be
- females arrested in their development—as is the case in social
Hymenoptera—and by supposing the soldiers to be males with —
arrested and diverted development.
The observations already made show that this is not the case.
It has been thoroughly well ascertained by Lespés and Fritz Miiller
that in various species of Calotermes the soldiers are both males
and females. Lespés and Grassi have shown that the workers of
Termes lucifugus are of male and female sex, and that this is
also true of the soldiers. Although the view of the duality of
the sexes of these forms was received at first with incredulity, it
appears to be beyond doubt correct. Grassi adds that in all the
individuals of the workers and soldiers of Z’ermes lucifugus the
sexual organs, either male or female, are present, and that they
are in the same stage of development whatever the age of the
individual. This statement of Grassi’s is of importance because
it seems to render improbable the view that the difference of form
of the soldier and worker arises from the arrest of the develop-
XVI TERMITIDAE 375
y
ment of their sexual organs at different periods. The fact that
sex has nothing whatever to do with the determination of the
form of workers and soldiers may be considered to be well
established. |
The statement that the young are all born alike is much
more difficult to substantiate. Bates said that the various forms
could be detected in the new-born. His statement was made,
however, merely from inspection of the nests of species about
which nothing was previously known, and as it is then very
difficult to decide that a specimen is newly hatched, it is probable
that all he meant was that the distinction of workers, soldiers,
and sexual forms existed in very small individuals—a statement
that is no doubt correct. Other observers agree that the young
are in appearance all alike when hatched, and Grassi reiterates his
statement to this effect. Hence it would appear that the differ-
ence of form we are discussing arises from some treatment subse-
quent to hatching. It may be suggested, notwithstanding the
fact that the young are apparently alike when hatched, that they
are not really so, but that there are recondite differences which are
in the course of development rendered conspicuous. This con-
clusion cannot at present be said with certainty to be out of the
question, but it is rendered highly improbable by the fact
ascertained by Grassi that a specimen that is already far advanced
on the road to being an ordinary winged individual can be diverted
from its evident destination and made into a soldier, the wings
that were partially developed in such a case being afterwards
more or less completely absorbed. This, as well as other facts
observed by Grassi, render it probable that the young are truly,
as well as apparently, born in a state undifferentiated except as
regards sex. Fig. 230 (p. 363) is designed to illustrate Grassi’s
view as to this modification; the individual A is already far
advanced in the direction of the winged form C, but can never-
theless be diverted by the Termites to form the adult soldier B.
According to the facts we have stated, neither heredity nor
‘sex nor arrest of development are the causes of the distinctions
between worker and soldier, though some arrest of development is
common to both; we are therefore obliged to attribute the dis-
tinction between them to other influences. Grassi has no
hesitation in attributing the anatomical distinctions that arise
between the soldiers, workers, and winged forms to alimentation.
376 NEUROPTERA CHAP.
‘Food, or the mode of feeding, or both combined, are, according
to the Italian naturalist, the source of all the distinctions,
except those- of sex, that we see in the forms of any one ispecies
of Termite. —
-Feeding.—Such knowledge as we possess of the food-habits
of Termitidae is chiefly due to Grassi; it is of the very greatest
importance, as giving a clue-to much that was previously obscure
in the Natural History of these extraordinary creatures.
In the abodes of the Termites, notwithstanding the enormous
numbers of individuals, cleanliness prevails; the mode by which
it is attained appears to be that of eating all refuse matter.
Hence the alimentary canal in Termitidae contains material of
various conditions of nutritiveness. These Insects eat their cast
skins and the dead bodies of individuals of the community ; even
the material that has passed through the alimentary canal is
eaten again, until, as we may presume, it has no further nutritive
power.. The matter is then used for the construction of their
habitations or galleries, or is carried to some unfrequented part
of the nest, or is voided by the workers outside of the nests;
the pellets of frass, 7.e alimentary rejectamenta, formed by —
the workers frequently betraying their presence in buildings
when none of the Insects themselves are to be seen. The
aliments of Calotermes flavicollis are stated by Grassi and
Sandias to be as follows: (1) wood; (2) material passed
from the posterior part of the alimentary canal or regurgi-
tated from the anterior part; (3) the matter shed during
the moults; (4) the bodies of other individuals; (5) the
secretion of their own salivary glands or that of their fellows;
(6) water. Of these the favourite food is the matter passed .
from the posterior part of the alimentary canal. We will speak of
this as proctodaeal food. When a Calotermes wishes food it strokes
the posterior part of another individual with the antennae and
palpi, and the creature thus solicited yields, if it can, some
proctodaeal food, which is then devoured. Yielding the proctodaeal
food is apparently a reflex action, as it can be induced by friction
and slight pressure of the abdomen with a small brush. The
material yielded by the anterior part of the alimentary canal may
be called stomodaeal product. It makes its appearance in the
mouth in the form of a microseopie globule that goes on in-
creasing in size till about one millimetre in diameter, when it is
TS aye es
XVI TERMITIDAE | 377
either used for building or as food for another individual. The
mode of eating the eédysial products has also been described by
Grassi and Sandias. When an individual is sick or disabled it
is frequently eaten alive. It would appear thatthe soldiers are
great agents in this latter event, and it should be noticed that
owing to their great heads and mandibles they can obtain food by
other means only with difficulty. Since they are scarcely able to
gnaw wood, or to obtain the proctodaeal and stomodaeal foods,
their condition may be considered to be that of permanent
hunger, only to be allayed by carnivorous proceedings. When
thrown into a condition of excitement the soldiers sometimes
exhibit a sort of Calotermiticidal. mania, destroying with a few
strokes five or six of their fellows. It is, however, only proper to
say that these strokes are made at random, the creature having
no eyes. The carnivorous propensities of Calotermes are ap-
parently limited to cannibalism, as they slaughter other white
ants (Termes lucifugus) but never eat them.
~The salivary food is white and of alkaline nature; when
excreted it makes its appearance on the upper lip. It is used
either by other individuals or by the specimen that- produced
it; in the latter case it is transferred to the lower lip and
swallowed by several visible efforts of deglutition. The aliments
we have mentioned are made use of to a greater or less extent by
all the individuals except the very young; these are nourished
only by saliva: they commence taking proctodaeal and stomo-
daeal food before they can eat triturated wood.
Royal Pairs.—The restriction of the reproductive powers of
a community to a single pair (or. to a very restricted number. of
individuals) occurs in all the: forms of social Insects, and in
all of them it is concomitant with a prolongation of the repro-
ductive‘ period far exceeding what is natural in Insects. We are
not in a position at present to say to what extent the lives of the
fertile females of Termitidae are prolonged, there being great: diffi-
culties in the way of observing these Insects for long periods owing
to their mode of life ; living, as they do, concealed from view, light
and disturbance appear to be prejudicial to them. We have every
reason to believe, however, that the prolongation extends as a
rule over several years, and that it is much greater than that of
the other individuals of the community, although the lives of
even these latter are longer than is usual in Insects; but this
378 NEUROPTERA CHAP.
point is not yet satisfactorily ascertained. As regards the males
there is reason to think that considerable variety as to longevity
prevails. But the belief is that the royal males of Termitidae also
form an exception to other Insects in the prolongation of the
terminal periods of their lives. In Hymenoptera, male in-
dividuals are profusely produced, but their lives are short, and
their sole duty is the continuation of the species by a single
| act. We have seen that
Grassi is of opinion that
a similar condition of
affairs exists at present
with Termes lucifugus in
Sicily, but with this ex-
ception it has always been
considered that the life
of the king Termite is,
roughly speaking, con-
temporaneous with that
of the queen; it is said
that in certain species the
king increases in bulk,
though not to an extent
that can be at all com-
pared with the queen.
It must be admitted.
that the duration of life
of the king has not been
Fic. 235.—Royal pair of Termes malayanus from sufficiently established, for
Singapore, taken out of royal cell. A, A, King, the coexistence of a king
intra and dora views BB. quem, donald with g queen in the
royal cell is not incon-
sistent with the life of the king being short, and with his replace-
ment by another. Much that is imaginary exists in the litera-
ture respecting Termites, and it is possible that the life of the
king may prove to be not so prolonged as has been assumed.
Returning to the subject of the limitation of the reproduction
of the community to a single pair, we may remark that a priori
one would suppose such a limitation to be excessively unfavour-
able to the continuation of the species; and as it nevertheless is
the fact that this feature is almost, if not quite, without exception
XVI TERMITIDAE 379
in Insect societies, we may conclude that it is for some reason
absolutely essential to Insect social life. It is true that there
are in Termitidae certain partial exceptions, and these are so
interesting that we may briefly note them. When a royal cell is
opened it usually contains but a single female and male, and
when a community in which royal cells are not used is inspected
it is usually found that here also there are present only a single
fertile female and a single king. Occasionally, however, it
‘happens that numerous females are present, and it has been
‘noticed that in such cases they are not fully matured females, but
are imperfect, the condition of the wings and the form of the
anterior parts of the body being that of adolescent, not adult
Insects. It will be recollected that the activity of a community
of Termites centres round the great fertility of the female ;
without her the whole community is, as Grassi graphically puts
it, orphaned; and the observations of the Italian naturalist
make it clear that these imperfect royalties are substitution
queens, derived from specimens that have not undergone the
natural development, but have been brought into use to meet the
calamity of orphanage of the community, The Termites appar-
ently have the power of either checking or stimulating the reproduc-
tive organs apart from other organs of the body, and they appear to
keep a certain number of individuals in such a condition that in case
of anything going wrong with the queen, the reserves may be brought
as soon as possible into a state of reproductive activity. The in-
dividuals that are in such a condition that they can become pseudo-
royalties are called complementary or reserve royalties, and when
actually brought into use they become substitution royalties. It
is not at present quite clear why the substitution royalties should be
in such excess of numbers as we have stated they were in the case
we have figured (Fig. 236), but it may be due to the fact that when
the power of the community is at a certain capacity for supporting
young a single substitution royalty would not supply the requisite
producing power, and consequently the community adopts a
greater number of the substitution forms. Termites are utterly
regardless of the individual lives of the members of the community,
and when the reproductive powers of the company of substitution
royalties become too great, then their number is reduced by the
effective method of killing and eating them.
According to Grassi’s observations, the communities of Termes
380 ‘daz NEUROPTERA CHAR.
lucifugus are now kept up in Sicily almost entirely by substitution
royalties ; the inference being
that the age of each com-
munity has gone beyond the
capacity for life of any single
royal queen.
The substitution- royal-
ties’ are, as we have said,
| ~ ealled neoteinic (véos, youth-
ful, reivw, to belong to), be-
cause, though they carry on
the functions of adult Insects,
they retain the juvenile con-
dition in certain respects,
and ultimately die without
having completed the normal
development. © The. pheno-
menon.is not quite peculiar
to Insects, but oecurs in
- Fig. 236.—Pair of neoteinic royalties, taken some other animals having
from the royal chamber of Termes mirabilis r Z
at Singapore by Mr. G. D. Haviland, The @ Well = marked’ ometqmios
queen was one of thirteen, all in a nearly phosis, notably in the Mexi-
similar state. A, king; B, C, queen. ean Axolotl:1
A point of great importance in connexion with the neoteinic
royalties is that they are not obtained from the instar im-
mediately preceding the adult state, but are made from Insects
in an earlier stage of development. The condition immediately
preceding the adult state is that of a nymph with long wing-
pads; ‘such specimens are not made into neoteinic royalties, but
nymphs of an earlier stage, or even larvae, are preferred. It. is
apparently by an interference with one of these earlier stages of
development that the “nymphs of the second form,” which have ~
for long been an enigma to zoologists, are produced.
Post-metamorphic Growth.—The increase of the fertility of
the royal female is accompanied by remarkable phenomena of
growth. Post-metamorphic growth is a phenomenon almost
unknown in Insect life, except in these Termitidae; distension
not infrequently occurs toa certain extent in other Insects, and
Veale
> Ct
1 Camerano,. Bull. Soc. ent. Ital. xvii. 1885, p- 89; and Kollmann, Verh. Ges.
Basel, vii. 1883, p. 391.
XVI | TERMITIDAE 381
is usually due to the growth of eggs inside the body, or to the
repletion of other parts. But in Termitidae there exists post-
metamorphic growth of an extensive and complex nature; this
growth does not affect the sclerites (i.e. the hard chitinous parts
of the exo-skeleton), which remain of the size they were when the
post-metamorphic growth commenced, and are consequently mere
islands in the distended abdomen (Fig. 236, B,C). | The growth
is chiefly due to a great increase in number and size of the egg-
tubes, but there is believed to be a correlative increase of various
other parts of the abdominal as distinguished from the anterior
regions of the body. A sketch of the distinctions existing
between a female of a species at the time of completion
of the metamorphosis and at the period of maximum fertility
does not appear to have been yet made,
New Communities.—The progress of knowledge in respect of
Termitidae is bringing to light a quite unexpected diversity of
_ habits and constitution. Hence it is premature to generalise on
important matters, but we may refer to certain points that
haye been ascertained in connexion with the formation of new
communities. The duration of particular communities and the
modes in which new ones are founded are still very obscure.
It was formerly considered that swarming took place in
order to increase the number of communities, and likewise for
promoting crossing between the individuals of different com-
munities. Grassi, however, finds as the result of his prolonged
observations.on Termes lucifugus that the swarms have no further
result than that the individuals composing them are eaten up.
And Fritz Miiller states* that in the case of the great majority
of forms known to him the founding of a colony by means
of a pair from a swarm would be just about as practicable
as to establish a new colony of human beings by placing a
couple of newly-born babes on an uninhabited island. It was
also thought that pairs, after swarming, re-entered the nests
and became royal couples. It does not, however, appear that
any one is able to produce evidence of such an occurrence. The
account given by Smeathman of the election of a royal couple
of Termes bellicosus is imperfect, as, indeed, has already been
pointed out by Hagen. It suggests, however, that a winged
pair after leaving the nest do again enter it to become king
1 Jena. Zeitschr. Naturw. vii. 1873, p. 458.
Ki de NEUROPTERA CHAP.
and queen. The huge edifices of this species described by
Smeathman are clearly the result of many years of labour, and at
present substitution royalties are not known to occur in them,
so that it is not improbable Smeathman may prove to be correct
even on this point, and that in the case of some species mature
individuals may re-enter the nest after swarming and may become
royal couples. _On the whole, however, it appears probable that
communities of long standing are kept up by the substitution
royalty system, and that new communities when established are
usually founded by a pair from a swarm, which at first are not
in that completely helpless condition to which they come when
they afterwards reach the state of so-called royalty. Grassi’s
‘observations as to the sources of food remove in fact one of the
difficulties that existed previously in regard to the founding of
new colonies, for we now know that a couple may possibly bear
with them a sufficient supply of proctodaeal and. stomodaeal
aliment to last them till workers are hatched to feed them, and
till soldiers are developed and the community gradually assumes
a complex condition. Professor Perez has recently obtained * the
early stages of a community from a winged pair after they had
been placed in captivity, unattended by workers. Miiller’s
observation, previously quoted, is no doubt correct in relation
to the complete helplessness of royal pairs after they have
been such for some time; but that helplessness is itself only
gradually acquired by the royal pair, who at first are. able to
shift for themselves, and produce a few workers without any
assistance,
Anomalous Forms.—Miiller has described a Calotermes under
the name of C. rugosus, which is interesting on account of the
peculiar form of the young larva, and of the changes by which
it subsequently becomes similar in form to other species of the
genus. We represent the development of this larva in Fig.
237. We may call attention to the fact that this figure illus-
trates the large size of the paunch, which is so extraordinary
in some of the states of the Termitidae. i
It will be recollected that the genus Calotermes is destitute
of workers. There is another genus, Anoplotermes, in which the
reverse condition prevails, and the soldier is absent; this is the
only case. yet known in which such a state of affairs exists.
1 CR. Ac. Paris, cxix. 1894, p. 804..
rT an an
XVI? TERMITIDAE ! 383
The species is called A. pacificus by Fritz Miller; it differs from
other Termitidae in possessing a proventriculus destitute of tritu-
rating ridges. The nests of this species are utilised by a little
Eutermes (E. inguilinus Miiller) for its own advantage; whether
by first ‘destroying the Anoplotermes or whether by merely taking
possession of the nests abandoned by their owners is not known.
It is a most remarkable fact |
that the Hutermes resembles
the | TERMITIDAE 385
of them is a patch over which the grass has been cut quite
short. Mr. Haviland followed these holes by digging for a
distance of 20 feet and to a depth of 54 feet; they remain
uniform in size except that near the aie there may
be one or two chambers in which the grass is temporarily
stored, but these do not hold more than would be collected
in an hour or two. As the burrow descends it is occa-
sionally joined by another, and at the point of junction there
is usually a considerable widening. Sometimes they run straight
for 6 or 7 feet, sometimes they curve abruptly, sometimes they are
nearly horizontal, but near the mouth may be almost vertical in
direction. These Termites are very local, but the specimens are
numerous when found. Mr. Haviland dug for these Insects at
two places on the Tugela river, one of them being at Colenso. It
is much to be regretted that he was unable to reach the nest.
We figure a soldier selected from specimens sent by Mr. Haviland
to the Cambridge University Museum. This Insect is apparently
much smaller than Smeathman’s 7. viarwm. Other species of
Termitidae have been described * as forming underground tunnels
in Africa, but none of the a have ‘yet been satisfactorily
identified.
It was stated by Smeathman that some species of Termites
had chambers in their habitations in which grew a kind of fungus
used by the Insects for food; Mr. Haviland is able to confirm
Smeathman in this particular; he having found fungus-chambers
in the nests of more than one species both in Binpation and
South Africa (Fig. 240). |
Habitations.—In nothing do Termites differ more than in
the habitations they form.. Sometimes, as we have mentioned in
the case of Calotermes, there is no real structure formed; only a
few barriers being erected in burrows or natural hollows in wood.
In: other cases very extensive structures are formed, so that the
work of the Termites becomes a conspicuous feature in the land-
scape. This is of course only the case in regions that are not
much interfered with by man; the great dwellings spoken of
by Smeathman and others soon disappear from the neighbourhood
of settlements, but in parts of Africa and in Australia large
dwellings are still formed by these creatures. In the latter part
of the world there exists a very remarkable one, formed by an
She tt pane. Dae - 1. Kolbe, Ent.-Nachr. xiii. 1887, p. 70. ,
VOL. V 2 C
386 NEUROPTERA CHAP,
undetermined species called by the officers and crew of her
Majesty’s ship Penguin the “compass ant.” The outline of one
of the structures formed by this Termite we represent in Fig. 239.
Mr. J. J. Walker, to whom we are indebted for the sketch from
which this figure is taken, has also favoured us with the following
extract from his diary, of date 4th August 1890: “The most in-
teresting feature in the scenery (about forty miles inland from Port
Darwin) was the constant succession of huge mounds raised by
the Termites, of which I had seen some comparatively small
examples in my rambles near Port Darwin; but these exceeded
in dimensions all I had ever seen. The most frequent as well as
the largest kind was usually of a reddish or ferruginous colour
outside, and generally almost cylindrical in shape with obtusely-
pointed top, but nearly always more or less weather-worn, with
great irregular buttresses and deep ruts down the sides; many of
them look like ruined towers in miniature. Their usual height
was from 8 to 10 feet, but many were much higher, and some
attained an (estimated) elevation of at least 20 feet: Another
kind, seen only in one or two places along the line, was of a much ~
more singular character; they averaged only 4 to 5 feet high,
were built of a dark-gray mud,
and in shape were like thin flat
wedges set upright (see Fig. 239),
reminding one of tombstones in
a churchyard. But the most ©
remarkable feature about these
mounds was that they had all
the same orientation, viz. with
the long faces of the wedge
; “iat B _— pointing nearly north and south.
Fi, 280. Termitariay of compass or ney Why this is so I am quite at a
face extending south and north ; B, cross- less to imagine, and I much re-
Serres eret that I had no opportunity
of closely examining these most singular structures. A third
kind of mound, usually not exceeding 2 feet in height, was of a
simple, acute, conical figure, and generally of a gray colour some-
what paler than the last, f.
The material used for the construction of the dwellings is
either earth, wood, or the excrement of the Termites. The huge
edifices mentioned by Smeathman are composed of earth cemented
ee a ae ae
= ae
XVI TERMITIDAE 387
together so as to look like stone or brick, and the buildings appear
to be almost as strong as if they were eeey ésnstrneted with
these materials. In many cases
the substance used is comminuted
wood that has passed one or
more times through the alimen-
tary canal of the Insects, and
may therefore be called excrement.
Whether the stone-lke material
is made from earth that has
passed through the alimentary :
canal or from grains gathered Mf a eee of Termitarium of
ermes angustatus, 8. Africa, showing
for the purpose has not been fungus chambers and orifices of com-
well ascertained. In any case = ™™™catiom |
the material is cemented together by means of the secretions of
glands. Dudley and Beaumont have described the process of
construction, in a species observed by them, saying that earth
is brought*and placed in position by the mandibles, and cemented
by liquid from the abdomen! Von Jhering says” that some
species form the exterior walls of their dwellings of stene-like
material, but make use of woody matter for the construction
of the interior. Smeathman has described the nest. of Temes
bellicosus. The whole of the very strong external wall consists
of clay-like material, cemented by the secretions of the Termites
to a very firm consistence. The royal cell is built of the same
material as the framework of the nest; whilst the nurseries
in which the young are chiefly found are built of woody
material, and are always covered with a kind of mould—the
mycelium of a fungus—and plentifully sprinkled with small
white bodies, which, under the microscope, are found to be filled
with a number of oblong, spore-like cells. 3
These nurseries rest on the clay-like framework of the nest,
but are not attached thereto; they in no way support it, or one
another, indeed they have the appearance of being constantly
added to on their upper margins and constantly eaten away on
their under parts. Fig. 240 represents the appearance of the
upper: boundary of a nursery taken from a nest of Zermes angus-
tatus. The small white bodies, mentioned above, have dis-
appeared: the mycelium of the fungus, though not shown in the
1 Trans. N. York Ac. viii. 1889, p. 91. 2 Congr. internat. Zool. ii, 1892, p. 249,
388 NEUROPTERA CHAP. _
figure, is still visible on the specimen from which it was drawn,
and.gives rise to a whitish, glaucous appearance.
In various parts of the world nests formed on trees by Termites
are to be seen; these tree nests are, it would appear, in some cases
only parts of a community, and are connected with the main body —
by galleries. In other cases nests are formed in various positions of ©
advantage; Messrs. Hubbard and Hagen have given us an account *
of some of these—probably the work of Hutermes ripperti—as seen
in Jamaica, They describe the nests as spherical or conical masses,
looking externally as if composed of loamy earth; they are placed
on trees, fences, or walls; they vary in size from that of a man’s
fist to that of a hogshead; they appear to be composed of finely
comminuted. wood fastened together by saliva. .These nests are
formed on the same principle as those of the wasps that make
nests hanging to trees and bushes, as they consist of an external
protecting envelope covering a comb-like mass in the interior.
At the bottom of the nest there is a covered gallery leading to
the earth, where the main nest appears to be situate; galleries
also are constructed so as to lead to the tops of trees and other
places, in such a manner that the Termite can still keep up its
peculiarity of working and travelling in tunnels and yet roam
over a large area; the activity of these Termites continues day
and night. _In each nest there is a queen, who lays eggs that
are removed by the worker Termites to. the bottom of the nest.
The young are fed on a prepared food, consisting apparently of —
comminuted vegetable matter, of which considerable masses are
laid in store. Some of the nests are rich in containing many
pounds’ weight of this material, while others are apparently quite
destitute of it. There is a soldier form and at least two kinds of -
workers. -. Some species of true ant frequently shares the nest of
these white ants, but on what terms the two kinds of Insects lve
together is not stated. |
Termite Ravages.—In countries whose climate is favourable
to their constitutions certain kinds of Termites become of great
importance to our own species. Owing to their taste for woody
matter and to their habit of working in concealment, it 1s no
‘uncommon thing for it to be discovered that Termites have
obtained access to a building and have practically destroyed the
wooden materials used in its construction ;. all the interior of the
- 4+. P, Boston. Soc. xix, 1878, p. 267.;.and xx. 1881, p. 121.
>
XVI TERMITIDAE 389
wood being eaten away and only a thin outer shell left intact.
A Termite, 7. tenuis, was introduced—in what manner is not
certainly known'—to the Island of St. Helena, and committed
such extensive ravages there that Jamestown, the capital, was
practically destroyed and new buildings had to be erected. Other
such cases are on record. Destructive species can sometimes be
destroyed by placing in the nests a portion of arsenicated food.
This is eaten by some individuals, who perish in consequence ;
and their dead bodies being consumed by their comrades, the colony
becomes checked if not exterminated.
The number of described species of Termitidae does not much
exceed 100, but this is certainly only a. small portion of those
existing, the total of which may probably reach 1000 species.
Termitidae are classed by some naturalists with the Orthoptera,
and they have a great deal in common with some of the cursorial
division of that Order, more particularly Forficulidae and Blattidae;
but they differ from Orthoptera in the nature and form of the
wings. They are also classed by some, with a few other forms, as
a separate Order of Pseudo-Neuroptera called Corrodentia, but this
is not a very satisfactory course, as the Termitidae do not agree
closely with the forms associated with them, while the aggregate
so formed is far from being very distinct from other forms of
Neuroptera. On tho whole the best plan appears to be to treat the
Termitidae as forming a distinct family of the Order Neuroptera,
or to make it a distinct Order, as proposed by Grassi. Packard
now associates Termites in an Order with the biting-lice, and
calls it Platyptera.
Fossil Termites.—Termitidae were very abundant in Rasthacry
times, and the genera appear to have been then much the same as
at present. In.Mesozoic strata the remains of true Termitidae
apparently exist in the Lias in Europe, but farther back than
this the family has not been satisfactorily traced. It was formerly
supposed that Termitidae existed in the Carboniferous strata, but
this appears to be very doubtful; and the fossil remains of that
epoch, which were presumed to be those of Termites, are now
referred by Scudder and others to the Neuropteroid division of the
Order Palaeodictyoptera, an Order which is formed entirely of
Palaeozoic fossil remains.
1 According to Melliss, it is thought that the Insect may have been carried to
the island in a captured slave-ship. Melliss, S#. Helena, 1875, p. 171.
CHAPTER XVII
NEUROPTERA CONTINUED—PSOCIDAE (BOOK-LICE AND DEATH-
WATCHES )—THE FIRST FAMILY OF AMPHIBIOUS NEUROPTERA
(PERLIDAE, STONE-FLIES).
Fam. IV. Psocidae—Book-Lice, Death-Watches.
Minute Insects with slender, thread-like, or hair-like antennae ;
four delicate membranous wings, the front pair of which are
the larger; their neuration is not
abundant and is irregular, so that
the cells are also irregularly ar-
ranged ; the transverse nervules are
only one or two in number Pro-
| thorax very small, in the winged
ia) eet tae Veikaiin forms quite concealed between the
England. (After M‘Lachlan.) head and the large mesothorax ; this
latter closely connected with, or fused
with, the metathorax. Species quite wingless, or with wings
unfitted for flight, exist ; in them the prothorax is not so ex-
tremely small, while the mesothoraxz is smaller than in the
winged forms. Tarsi of two or three segments. Metamorphosis
slight, marked chiefly by the development of wings and ocelli.
THE Psocidae are without exception small and soft-bodied Insects,
and are only known to those who are not entomologists by the
wingless forms that run about in uninhabited or quiet apart-
ments, and are called dust-lice or book-lice. They are perhaps
more similar to Termitidae than to any other Insects, but
the two families differ much in the structure of their wings, and
are totally dissimilar in the nature of their lives.
1 In some exotic species there is a dense network on a part of the anterior wing.
CHAP. XVII ° PSOCIDAE . 391
The antennae consist of eleven to twenty-five joints, or even
more, about thirteen being the
usual number; the basal two are
thicker than the others, and are
hy destitute of setae or pubescence
3 such as the others possess. The
-. maxillae and labium are remark-
able. The former possesses a
peculiar hard pick or elongate
rod; this is considered by many
naturalists to be the inner loke, Fic. 242.—Transverse horizontal section
but Burgess thinks it more prob- peteees: Le a ae
ably an independent organ,' as it =P, stipes; m.m, muscles ; m.s, socket
APSE Te ‘ of mandible,
has no articulation of any kind
with the outer lobe. The latter is remarkably thick and fleshy ;
‘ the palpus is 5-jointed. Other
| ~ authorities consider the pick to
be certainly the inner lobe; if’
it be not, the latter is quite
wanting. Hagen agrees with
Burgess in stating that the
pick slides in the outer lobe as
in a sheath, The labium has
a large mentum and a ligula
divided anteriorly into two lobes;
at each outer angle in front
there is a globular projection,
which is doubtless the labial
palpus; reposing on the labium
there is a. large free lingua.
The labrum is large, attached
to a distinct clypeus, behind
Fic. 243.—A, Front of head of Psocushetero- which there is a remarkable
morphus ; cl, post clypeus ; g, epicran- , .
ium: B, transverse horizontal section post-clypeus, which is usually
of post-clypeus of Psocus: cl, post- prominent as if inflated; to its
clypeus ; ¢.m, clypeal muscles; g, epi- ~
cranium ;¢, tendons ;/.m,labial musclein Inner face are attached several
section ; 0e, oesophagus ; 0e.b, oesopha- _
geal bone. (After Burgess and Bertkau. ) muscles which Baro shes te to be
inserted on a plate placed below
the anterior part of the oesophagus, and called by Burgess the
1 P. Boston Soc. xix. 1878, p. 292.
392 NEUROPTERA CHAP,
oesophageal bone; under or within the lingua there is a pair
of lingual glands, Judging from Grosse’s. study of the mouth
of Mallophaga, we may conclude that the oésophageal bone will
prove to be a sclerite of the hypopharynx. , The eyes of the
winged forms are frequently remarkably convex, and there are
also three ocelli, triangularly placed on the vertex. The head
is free and very mobile. The coxae are rather small, exserted,
contiguous; the sterna small. _ The abdomen has usually, ten
segments, though sometimes only nine can be detected. -
The thorax in Psocidae usually: looks as if it consisted of
only two segments. This is due to’ two opposite conditions: (1)
that in the winged forms the prothorax is reduced to a plate
concealed in he. fissure between the head. and the mesothorax
bearing the first pair of wings; (2) that in the wingless forms
(Fig. 24.7 ), though the Srotheite is distinct, the meso- and meta-
thorax are fused into one segment.
The internal anatomy is only very incompletely known.
Nitzsch * has, however, described the alimentary canal and the
reproductive organs of Clothilla pulsatoria. The former is |
Fic. 244.—Reproduc-
tive organs of Clo-
thilla pulsatoria.
A, Male; a, vesi-
culae seminales ; 0,
testes ; ¢, vasa de-
ferentia; d, ejacu-
latory duct. B,
Female ; a, 0, egg-
tubes ; ¢, oviduct ;
d, uterus, contain-
ing egg; @, acces-
sory gland (the en-
veloping sac in sec-
tion) ; f, its duct.
(After Nitzsch.)
remarkably sunple: no proventriculus or crop was found; the
stomach is very elongate, and consists of a sac-like anteriae
portion and an elongate, tubular posterior part. There are four
Malpighian tubes. The posterior part of the canal is remark-
ably short, the small intestine being scarcely as long as the
rectum. The ovaries (Fig. 244, B) consist of five egg-tubes on
each side; connected with the oviduct there is a peculiar acces-
sory gland consisting of a sac containing other small sacs each
? Germar, Mag. Entomol. iv. 1821, p. 276, pl. ii.
XVII | -PSOCIDAE 393
with an elongate efferent duct; the number of the secondary
sacs varies from one to four according to the individual. The
testis (Fig. 244, A, b) is a simple capsule; connected with the
base of the ejaculatory duct there is a pair of elongate accessory
glands or vesiculae seminales,
The life-history has never been satisfactorily sketched. The
young greatly resemble the old, but have no ocelli or wings, and
sometimes the tarsi are of two joints, while in the adult they
have three. The antennae have also in these cases a less number
of joints in the young stage. The food is animal or vegetable
refuse substances; many live on fungoid matter of various kinds,
mouldy chaff being, it is said, a favourite pabulum; the mould
on palings is a source of food to many; others live on the -rust-
fungi of leaves, and many frequent the bark of trees. They are
able to spin webs, probably by the aid of the lingual glands;
the eggs are deposited, in some cases, on leaves and covered. with
a web. Hagen says that a peculiar organ, possibly a gland—he
ealls it a Sead? —exists at the base of the tarsal claws. In’ our
climate most. of the species pass the winter in the egg-state.
There may be two generations in a year, perhaps more.
The nomenclature of the wing-veins of Psocidae has given
rise to much discussion.” The
system shown in the accom-
panying figure is probably the
most convenient ; the subcos-
tal vein (2) is always obscure,
and sometimes can only be
detected by very minute @X- Fic. 245.—Anterior wing of Elipsocus brevi-
amination. Some interesting vu (Afte: Reuter) 1, Costa vein
information as to the minute branches of cubitus; 5, sector of the radius ;
structure and mode of forma- — °™ 10*S thereof.
tion of the wings and their nervures has been given by Hagen.°
In the young the wings first appear as buds, or outgrowths
of the sides of the meso- and meta-thorax ; afterwards eek pro-
thorax decreases, while the other two thoracic segments and
the wing-rudiments attached to them increase. The wings
from their very origin appear to be different from. those of the
Orthoptera, and the changes that take place in the thoracic
_ Psyche, iii. 1881, p..196. 2 Kolbe, Stettin. ent. Zeit. xli. 1880, p. 179.
3 Op. cit. p. 209, ete.
394 : NEUROPTERA CHAP.
segments in the course of the development, differ from those
that occur in Orthopiera. |
There are several peculiarities connected with the wings.
Frequently they exist,
though of no-~ use for
flight ; some Psocidae that
have __ perfectly - forme
wings are so reluctant to
dP. ty use them that, M‘Lachlan
ill “= says, they will allow them-
Fie. 246.—Micropterous form of Mesopsocus unt- selves to be crushed with-
punctatus. a, a, Wings. (After Bertkau.)
out seeking to escape by
flight. At certain periods, however, some Peocidae float. on
the wing in considerable numbers, especially in a moist still
atmosphere, and then drift about like the winged Aphididae,
which are frequently found with them. There is evidence
that individuals,’or generations, of some of the winged species
occur with only rudimentary wings; although this has been
denied by Kolbe, there can be no doubt about it. The form
figured above (Fig. 246) was described by Bertkau’ as a dis-
tinct genus, but was afterwards recognised by him? to be a
short-winged form of Mesopsocus unipunctatus. It is probable
that the adult female of this species has the wings always
micropterous, while the male has these organs of the full size.
In other species the condition of the rudimentary wings seems to
be quite constant. The facts concerning the wings of Psocidae
are so peculiar that Kolbe came to the conclusion that the
organs exist not because they are of use for flight, so much as
because it is the nature of an Insect to develop wings.”
Some of the species of Psocidae have never any trace of wings.
These apterous forms are mostly included in the division
Atropinae, and are usually very minute; it has been again and
again erroneously stated that they are the young state of winged
forms. Hagen kept a large colony of Atropos divinatoria for
some years in confinement, so that he saw numerous generations
as well as many specimens. He found the apterous condition
quite constant. |
1 Arch. f. Naturg. xlix. i. 1883, p. 99.
2 Verh. Ver. Rheinland, xxxix. 1882, Corr. -bl. p. 128.
3 Berlin ent. Zeit. xxviii. 1884, p. 36.
XVII , PSOCIDAE 395
The association of ocelli with wings is nearly constant in
Psocidae. The genus Clothilla—allied to -Atropos—possesses
very rudimentary wings but no ocelli. Hagen, however, found *
that in a certain locality no less than 12 per cent of the indi-
viduals of this species were provided with ocelliima most extra-
ordinary variation.
In some of these apterous forms there is found on each side
of the prothorax a tubercular prominence which, according to
Hagen, can be considered only as the rudiment of a wing that
never develops. Though no existing Insect is known to possess
rudimentary wings on the prothorax, we have previously men-
tioned (p. 344) that in the Carboniferous epoch appendages of
the nature alluded to were not very rare.
A genus of living forms—AHyperetes—in which the three
thoracic segments are well developed, but in which there are no
alar appendages or rudiments, is considered by Hagen to be more
primitive than the Psocidae found in amber to which we shall
subsequently allude.
The number of described species of Psocidae does not reach
two hundred; we have, however, thirty species or more in
Britain.” Nietner observed about the same number in the
immediate vicinity of his house in Ceylon. The isolated and
remote Hawaiian group of islands is remarkably rich in Psocidae.
Two thousand is a moderate estimate of the number of existing
species. The largest forms yet discovered belong to the Brazilian
genus Thyrsophorus; they attain, however, a breadth of only
about one inch with the wings fully expanded. The Cuban
genus Hmbidopsocus is said to be of great interest from its
approximation to Embidae. It is at present very inadequately
known. .
One (or more) very minute Insects of this family—Clothilla
pulsatoria according to Hagen, Atropos*® divinatoria according to
some other authors—is widely known under the name of the
death-watch, owing to its being believed to make a peculiar
1 Stettin. ent. Zeit. xliv. 1883, pp. 299, 305.
2 For the British species, see M‘Lachlan, Ent. Month. Mag. iii. 1867, p. 177.
3 The genera Atropos and Clothilla were named after the two fates Atropos and
Clotho. Westwood attempted some years ago to complete the trio by establishing a
genus Lachesilla. This proved a failure, the genus being a misconception. As the
name Lachesis is in use in various branches of zoology, the desired circle of Psocid
fates is likely to remain always incomplete.
396 NEUROPTERA | CHAP,
ticking noise, supposed to be prophetic of the decease of some
individual—a human being we fancy,
not a death-watch. It is difficult
to believe that so minute and soft an
Insect can produce a sound audible to
human ears, and many entomologists are
of opinion that the sound in question
is really produced by a beetle—of the
genus Anobiwum—which lives in wood,
and that as the beetle may be concealed
in a hole, while the Clothilla is seen
running about, the sound is naturally,
: though erroneously, attributed to the
Fic. 247.—A, Atropos divina- latter. But the rapping of the Ano-
ate f Mee biwm is well known, is produced while
the Insect is at large, and is said to be
a different noise from that of the Psocid; evidence too has been
given as to the production of the sound in a .workbox when the
Psocid was certainly present, and the most careful search failed
to reveal any beetle.
The Rev. W. Derham, who two hundred years ago was Rector
of Upminster, in Essex, and was well known as a distinguished
writer and philosopher, gave an account of the ticking of death-
watches to the Royal Society.’ This gentleman was a most
accurate and minute observer; he was well acquainted’ with the
ticking of the greater death-watch — Anobiwm — which he
describes very accurately, as well as the acts accompanying it,
the details he mentions being exactly such as occur at the present
time. He not only heard the ticking of the Psocid or lesser
death-watch, but repeatedly witnessed it. He says: “I am now
so used to, and skilful in the matter as to be able to see, and
show them, beating almost when I please, by having a paper
with some of them in it conveniently placed and imitating their
pulsation, which they will readily answer.” He also states that
he could only hear them beating when it was done on paper, and
that this death-watch will tick for some hours together without
intermission, with intervals between the beats, so that it much
resembles the ticking of a watch. The act of ticking was accom-
1 Phil. Trans. xxii. 1701, pp. 832-834 ; and xxiv. 1704, pp. 1586-1594, Plate 291,
Figs. 4, 5 (pp. 1565 to 1604 occur twice in this sacs
I i el
Lill
i
XVII PSOCIDAE 397
panied by rapping the front of the head on the paper, but Mr.
Derham could not be sure that the sound
was produced in that manner, because each
stroke was also accompanied by a peculiar
shudder, or recoil. After a prolonged tick-
ing he observed that another individual of
the other sex made its appearance. The
species figured by Mr. Derham more.
resembles a Hyperetes than it does either
of our two known book-lice, Atropos and yg. 948 The lesser death-
Clothilla. watch of Upminster.
: ‘ After Derham. mag-
Numerous species of Psocidae are pre- pee B, sea eh :
served in amber; Hagen’ has made a )
careful study, based on a considerable number of specimens,
of about thirteen such species. They belong to no less than
nine genera and five sub-families. Sphaeropsocus is the most
remarkable; this Insect. has a well-developed prothorax, as
is the case in the wingless
Psocids, and a pair of large
Wings or tegmitia meeting
by a straight suture along
+ the back, as is usual in
~"\ beetles, though quite un-
3 known in existing Psocidae.
Another species, Amphiento-
mum paradoxum, has the
body and appendages covered
with scales like a butterfly
or moth; other species, found
Fic. 249.—Sphaeropsocus kunowii. From jy oyum-copal or still liv-
amber. x30. (After Hagen.) 8 P
ing, have scales on various
parts of the body, but not to so great an extent as this amber
species. The genus Amphientomum is still represented in Ceylon
and elsewhere by living forms; Packard has figured some of the
‘scales ;? they appear to be extremely similar to those of Lepi-
‘doptera or Thysanura. The facts connected with this fauna of
‘amber Psocidae would seem to show that the family was formerly
more extensive and important than it is at present; we should
therefore expect to find numerous fossil forms in strata of date
1 Stettin. ent. Zeit.-xliii. 1882, p. 265: - ? P. Boston Soc. xiii. 1871, p. 407.
398 NEUROPTERA CHAP.
anterior to that of the amber; but this is not the case, all that:
is known as to fossil Psocidae being that Scudder has recently
ascribed traces of an Insect found in the Tertiary rocks of Utah
to this family as a distinct genus. ©
Fam. V. Perlidae.
Insects of moderate or large size, furnished with four membranous
wings ; these are usually complealy reticulate ; the hind pair
are much the larger, and have a large anal area of more
simple venation, which becomes plicate when folded. The
coxae are small, the legs widely separated. The larvae are
aquatic in habits ; the metamorphosis is slight.
SS SS
Ro So
SA
Fig. 250.—Pteronarcys frigida, mele. (After Gerstaecker.)
The Perlidae form a small family of Insects unattractive in their
general appearance. The life-history of each individual consists
of two abruptly contrasted portions; the earlier stage being
entirely aquatic, the later aerial. Hence the Perlidae come into
the amphibious division of Neuroptera. The definition we have
given above would, except as regards the texture of the front
wings and the aquatic habits of the larvae, apply to many
Insects of the Order Orthoptera. The Phryganeidae, another
XVII PERLIDAE 399
family of Neuroptera, have aquatic larvae and wings somewhat
similar in form to those of the Perlidae, but the members of the two
families cannot be confounded, as the Phryganeidae have hairy
front wings and large and contiguous coxae.
The antennae of the Perlidae are long, very flexible, and com-
‘posed of a very large number of joints. The parts of the mouth
vary a good deal. The mandibles and maxillae are usually rather
small, and all the parts of the mouth are of feeble consistence or
even membranous; the maxillary palpi are, however, well developed
and exserted from the mouth, five-jointed. The labium is short
and but little conspicuous. The mandibles in some forms are
almost membranous, but in other genera they are firmer and are
toothed. The labium is composed of a very large mentum, beyond
which is a large piece, usually undivided, bearing the four terminal
lobes; the three-jointed palpus ‘is seated on the side of the large
middle sclerite, which is no doubt of composite nature. Con-
siderable variety as to the lower lip prevails. The head is broad
and flat; there is an indistinctly-indicated clypeus, three—
more rarely two—ocelli, and on each side an eye neither very
large nor perfect. The prothorax is free, and has a flat,
margined notum. The meso- and the meta-thorax are large,
equal segments. The pro-, meso-, and meta-sternum are large
pieces; between the first and second, and between the second
and third there is an intervening membrane. The metasternum
is much prolonged backwards, and has on each side a_ peculiar
slit; similar orifices exist on the other sterna (Fig. 254, 0).
Newport, who has examined them in Pteronarcys, says that they
are blind invaginations of the integument; he calls them the
sternal or furcal orifices." According to this naturalist these very
_ peculiar openings pass into the body “as strong bone-like tubes,
diverging from the axis to the periphery of the body in the
immediate vicinity of some of the principal tracheae, but that
they do not in any way communicate with them, as they terminate _
abruptly as caecal structures.’ He thinks them analogous with
the endo-skeleton of other Insects; a view which cannot be con-
sidered sufficiently established. Laboulbéne states? that when
Perla parisina is seized and placed on its back, it does not move,
but emits a liquid at the base of the articulation of the legs.
WTr, Linn. Soc. xx. 1851, p. 433.
2 Bull, Soc. ent. France (4), viii. 1868, p. xxxvii.
400 |
NEUROPTERA CHAP.
This suggests that it may come from these sternal orifices. The
abdomen consists of ten dorsal plates, the
first being short, and of nine ventral;
the dorsal plates are much more ample
transversely than the ventral. Frequently
the hind body is terminated by two long;
many-jointed cerci, looking like antennae.
The coxae are small, not prominent, and
are directed outwards. The legs are
Slender, the tibiae often grooved. The
tarsi are three-joiited, terminating in two
claws and a more or less distinct pad.
In the genus Jsopteryx an auditory organ
has been described as existing in the legs,
in a position similar to that of the analo-
gous structures
Fig. 251.—Perla maxima. in ‘Termitidae
ha Say and Blattidae.
The.wings when closed repose flat on the:
back, and fold and overlap so that only
one is seen (Fig. 251); in this state
the costal portion of each front wing is ~
turned downwards, so as. to protect to
some extent, the sides. of the body.
The early stages are known, but have
not been described minutely, and there
appears to be very little information as
to the youngest life. All the species
are, when immature, aquatic in their
habits; the larvae greatly resemble the
perfect: Insects in form, though differing
in not possessing wings and in the
ocelli being merely opaque spaces.
They have rather large compound eyes ;
the future wings are represented by
lobe - like prolongations — varying in
length according to age—of the meso-
and-meta-notum., .In the Nemourae the
cerci are absent in the imago though
é
Zz
_ 4, .
j
Z Fe!
A:
0
*
f R
Vs, *
# \\
fy
nd »
Fig. 252. — Perla SP. nymph,
showing tracheal gills, Py: ré-
nées orientales. -
present in the young. The larvae of Perlidae are carnivorous
XVII PERLIDAE 401
and are able to swim well, the legs being provided with abundant
swimming hairs; they, however, as a rule, prefer to walk at the
bottom of the pool. or on rocks or boulders in the water they
live in. |
One of the most peculiar features of the Perlidae is their
respiratory system. Unfortunately the greatest differences of
opinion have prevailed on various matters in connexion with this
subject, and there are several points about which it is not possible
at present to express a decided opinion.
The larvae have no stigmata; it appears to be generally
agreed that there is ii them no means
of admitting air to the tracheal system
by means of orifices. Some breathe
entirely through the integument, the pro-
cess being aided by the accumulation of
tracheae at the spots where the breath-
ing orifices should be, and where the
integument is more delicate. Others,
however, possess gills in the form of pro-
truded bunches of filaments, connected
with tracheae in the manner shown in
Fig. 253. These filamentous branchiae 5,, 953 mvacheal gill and
occur in numerous species of the family, portion of a trachea of Ptero-
and are situate on various parts of the — ™7¥% (After Newport.)
body, but many species are destitute of them in genera, other mem-
bers of which possess the filaments. In some Nemourae instead
of bunches of filaments there are tubular projections on the pro-
thoracic segment; and in Dictyopteryx signata similar structures
oceur even in the cephalic region, Hagen stating * that there exists
a pair on the submentum and another on the membrane between
the head and the thorax. In the imago state, stigmata are present
in the normal fashion, there being two thoracic and six abdominal
pairs. In several species the filaments persist in the imago, so that
in these cases we meet with the curious condition of the coexistence
of branchiae with a well-developed and functionally active system
of spiracles ; this is the more curious because the creatures usually
have then nothing to do with the water, it having been ascer-
tained that in these cases the species live out of the water as other
terrestrial and aerial Insects do. These instances of persistence
; a 1 Zool. Anz. iii. 1880, p. 304.
VOL. V 2D
402 . NEUROPTERA CHAP,
of branchiae during the aerial life have been the source of some
perplexity ; the condition was shown to exist
in Pteronarcys by Newport, and has since
been demonstrated in various other forms.
Newport believed that the imago of Ptero-
narcys breathes by means of the gills,
although it lives out of the water and
possesses spiracles; and he informs us that
Mr. Barnston observed the Insect when on
the wing “constantly dipping on the surface
of the water.” Hence Newport concluded
that Pteronarcys in the winged state is “an
amphibious animal.” That a winged Insect
should live in the air and yet breathe by
means of gills would be truly extraordinary,
and there can be little doubt that Newport’s
CWI idea was erroneous. Hagen’ was able to
Fie, 54D ide of examine living imagos of the species in ques-
regalis,imago. (After tion. He found that they avoided the
Newport) ot water, and though he placed some indi-
gills; 0, sternal : :
orifices. viduals therein, yet they did not use the
gills. He also informs us that the branchiae
have, during life, a shrivelled appearance, indicating that they
are not functionally active, but are merely useless organs carried
over to the imago from the previous instar, in which they were
truly the means of obtaining air. Hagen also ascertained that
the spiracles of the imago are in a normal state, being adapted
for breathing, even as far back as the seventh abdominal
segment.
Great difference of opinion has prevailed as to the relations
of the branchiae to the stigmata, it having been contended that
the falling off of some of the branchiae left the stigmatic orifices.
The facts appear to be only consistent with the conclusion that
the two are totally independent organs. This subject has been
investigated by Palmén,’ who finds that in Perlidae—contrary
to what occurs in may-flies—the species are either entirely
destitute of gills, or these organs are persistent throughout
life. It is not to be inferred from this that the gills in. the
1 Stettin. ent. Zeit. xxxviii. 1877, p. 487.
2 Morphologie des Tracheensystems, Helsingfors, 1877, p. 21.
/
i hicsintnd
_—
XVII PERLIDAE
403
the exceptional Pteronarcys :
moult the gills usually become very
much contracted and concealed by the
new integument; in some cases they
merely appear as slight prominences
in the neighbourhood of the stigmata.
Pictet, Dufour, Newport, and Imhof?
have studied the internal anatomy. The
alimentary canal is remarkable for the
enormous oesophagus; there is no dis-
tinction between this and the crop. A
proventriculus is quite absent, and there
are no chitinous folds in the position it
usually occupies. The true stomach is
small, and only commences in the fourth
abdominal segment. It has a prolonged
lobe on each side in front, and in
addition to this eight sacs; thus there
are formed ten diverticula, fastened to
the posterior part of the oesophagus by
ligaments. The terminal portion of the
stomach is small, and apparently only
distinguished from the short intestine
by the point of insertion of the Mal-
pighian tubes; these vary in number
from about twenty to sixty. There are
two pairs of large salivary glands. In
Pteronarcys the caecal diverticula of the
stomach are wanting. In some Perlidae
the terminal parts of the gut are more
New-
complex than in Perla maxima ;
‘port figures both an ilium and colon
very strongly differentiated, and states
that these parts differ much in Perla and Pteronarcys.
perennibranchiate Perlidae are as conspicuous as they ave in
for it appears that at the final
fae ii
i)
(WD)
Fig. 255.—Alimentary canal and
“outline of body of Perla
maxima. (After Imhof.) 7,
Upper lip ; mh, buccal cavity ;
ap, common termination of
salivary ducts ; 0, oeso-
phagus ; s, salivary glands ;
ag, duct of salivary gland ;
6, anterior diverticula of
stomach ; dg, their ligaments
of attachment ; mp, Malpi-
ghian tubes; 7, rectum; af,
anal orifice.
Accord-
ing to him the stomach is embraced by a network of tracheae,
and Imhof tells us that he found the stomach to contain only air.
The brain is small, but, according to Imhof, consists of four .
amalgamated divisions; the infra-oesophageal ganglion is small,
1 Beitr. Anat. Perla maxima.
— Inaug.-Diss. Aarau, 1881.
404 NEUROPTERA CHAP. .
and placed very near the brain. There are three thoracic and
six abdominal ganglia on the
ventral chain. The nerves
to the wings are connected
with the longitudinal com-
missures of the ventral: chain
by peculiar, obliquely-placed,
short commissures. The repro-
ductive glands are peculiar,
inasmuch as in each sex
the pair of principal glands
is connected together in the
middle. The testes thus form
an arch consisting of a large
number of sub-spherical or
pear-shaped follicles; the
vasa deferentia are short in
Perla maxima, and there are
Fie, 250,—ahe yar of nite ovaries of Perl no vesiculae seminales; the
ceptaculum seminis concealing the orifice of ejaculatory duct is divided
the duct and an accessory gland. irita thee parts by coratrie
tions. In Pteronarcys and in Perla bicaudata, according to
Newport and. Dufour, the vasa deferentia are very long and
tortuous, and thers are elongate vesiculae
seminales. The arrangement of the ex-
tremely numerous egg-tubes is analogous to
that of the follicles of the testes, so that, as
Dufour says, there is but a single ovary ;
connected with the short, unpaired portion of
the oviduct, there is a large receptaculum
seminis, and near the terminal orifice of
the duct there is in P. maxima an eight- |
lobed accessory gland. Fic. 257.—Ege of Perla
The eggs are produced by Perlidae in ‘™awima. (After Imhof.)
c, chorion; d, oolemn ;
enormous numbers: they are rather small, gs, glass-like covering of .
but peculiar in form, and possess at one pag Feige 2
extremity a micropyle apparatus, covered canals penetrating
by a glassy substance through which Imhof = “°™"
could find no orifice. On the other hand, the chorion on another
part of the egg is perforated by several canals.
PEL
WIG I 47
BAK OMY |
Mid Vi i
4 Wy Y
3 wie ge
w yy y ‘wy
NAL BV fA
Y IU ee J
“xv PERLIDAE AOS
The Perlidae being of aquatic habits in their early stages,
and, notwithstanding their ample wings, very poor adepts in the
art of flying, are rarely found at any considerable distance from
their native element. They are specially fond of running water,
and delight in the neighbourhood of waterfalls, or other spots
where the current is broken by obstacles so that a foaming water
results. It is probable that the larvae which breathe by means
of gills find an advantage in living in strongly-aerated water.
Mountain streams and torrents are therefore specially affected by
them; but Pictet informs us that they do not like the waters
descending from glaciers. The food of the larvae is believed to
be chiefly young may-flies, or other small, soft creatures, and it
may possibly be owing to the absence of these that the Perlidae
do not affect the glacier streams. Although Perlidae are remark-
able for their capacity for enduring cold, it is possible that they
may require warmth of the water at some period of their
development, and this the glacier-streams cannot offer to them.
They are among the earliest Insects to appear in the spring in
Europe. Mr. Barnston says that on the Albany river in Canada
the nymph of Capnia vernalis comes up frequently in the cracks
of the ice and casts its skin there; “it frequently comes up
when the thermometer stands at freezing.” Of Nemoura glacialis,
which inhabits similar localities, he says that “it appears in the
spring (end of March or beginning of April) when the ice
becomes honeycombed, and even before then, at the same time
as Capnia vernalis. It pairs in the crevices of decaying ice.
The male has long antennae, and his wings are generally rumpled
as if glued together.” Newport entertained the idea that those
Perlidae that live at low temperatures are of lower organisation
than the other forms of the family.
It is a remarkable fact that several Perlidae frequently
have—like Nemoura glacialis—the wings of the male much
reduced in size; this being the contrary of the rule that
usually prevails among Insects to the effect that, when there is a
difference in the powers of flight, or even in the size of the wings,
it is the male that is superior. Mr. J. J. Lister met with a very
interesting Perlid at Loch Tanna in Arran at the beginning of
April 1892. In this Insect, which is, according to Mr.
M‘Lachlan, a form of Jsogenus nubecula, the wings of the female
(Fig. 258, B) are reduced to a size much less than those of ordinary
406 NEUROPTERA CHAP.
Perlidae, while those of the male (Fig. 258, A) are mere useless
rudiments, Morton has pointed out that in Scotland more
than one species of Zaeniopteryx occasionally produces. micro- _
pterous males, and he associates this phenomenon with the early
time of their appearance
“almost in winter.” In
Nemoura trifasciata this
reduction of the wings
takes another but equally
curious form; the hind
wings of the male being
long enough to cover the
body, while the anterior
pair are reduced to mere
rudiments.
The phenomena of mi-
Fic. 258.—Tsogenus nubecula, Loch Tanna. A, cropterism in Perlidae are
Male ; A’, wings of male more magnified; B, well worthy of more de-
wings of female. , : : .
tailed investigation. Mr.
Morton informs the writer that the male of Perla maxima (Fig.
251) in North Britain has the wings so short that they cannot
be of any use as organs of flight. In Central Europe the wings are
ample, as shown in our figure. In Perla cephalotes the male is
short-winged in both Britain and Central Europe; of the male of |
Dictyopteryx microcephala only the micropterous form is known to —
exist. In Jsogenus nubecula (Fig. 258) it appears that the
wings of the female are always more ample than those of the
male of the same locality, and that local micropterism affects
the. two sexes unequally. Within the Arctic circle this Insect
is usually of the Scotch form, though the male there occasionally
has more ample wings.
It has been observed that in some Perlidae the eggs, after
they have been extruded, are carried about by the female; for
what reason is not at all known. They are said to be enclosed
in a membranous capsule at the apex of the abdomen. The
number of eggs deposited is sometimes very large, amounting to
five or six thousand, and they are often of very minute size.
About twenty-four species of Perlidae occur in Britain.” The
1 Entom. Month. Mag. xxix. 1893, p. 249.
*No satisfactory systematic work of a general character on British Perlidae
XVII | PERLIDAE ' 407
species from all parts of the world existing in collections probably
scarcely exceed two hundred. The insignificance of this number
is no doubt chiefly due to the fact that these unattractive Insects
are rarely captured by collectors, and are so fragile that unless
good care is taken of them, specimens soon go to destruction
after being dried. Perlidae are known to occur in most parts
of the world, so that the number of species really existing may
reach two or three thousand. They are known to anglers as
stone-flies and creepers and are a favourite bait for trout.
The family in its character comes near to the Orthoptera,
especially to the more simple forms of Phasmidae, but the two
groups differ in the texture of the
front wings and in the structure
of the mouth-parts, as well as in
the different proportions of the
mesothorax and metathorax. Ac-
cording to Pictet,in the Australian
genus Lusthenia the trophi (Fig.
259) approach nearer to those of
the Orthoptera, so that it appears
possible that a more intimate con-
nexion will be found to exist as more Fic. 259.—A, Maxilla, B, labium
forms are discovered. Of the groups of usthenia spectabilis. (After
we include in Neuroptera, Perlidae Pte)
are in structure most allied to Sialidae, but the development in
the two groups exhibits very important distinctions. Brauer
treats the Perlidae as forming a distinct Order called Plecop-
tera, a name applied to the family by Burmeister many years
ago. |
Several species of Perlidae, considered to belong to existing
genera, have been found in amber. A _ fossil from the Eocene
deposits in the Isle of Wight and another from the Miocene of
Continental Europe are referred to the family. Brauer has
recently described’ some fossils from the Jurassic formation in
East Siberia as forming three genera, now extinct, of Perlidae.
Brongniart informs us” that several fossils have been found
exists. References to the scattered descriptions and notes will be found in the
Catalogue of British Neuroptera published by Entom. Soc. London, 1870.
1 Mem. Ac. Pétersb. (7) xxxvi. No. 15, 1889.
2 Insectes fossiles, etc., p. 407, 1893.
ee ST a nee > res a ee 1
408). Whit, NEUROPTERA ssid gs he
in the Carboniferous strata of Commentry that intr us
asserting that allies of Perlidae then existed. He considers: i
Carboniferous Insects to have belonged to a separate famil
Protoperlides. The fragments are, however, so small the
must await further information before forming a definite opini
as to these Protoperlides. < 3 eae a
CHAPTER XVIII
AMPHIBIOUS NEUROPTERA CONTINUED—ODONATA, DRAGON-FLIES
Fam. VI. Odonata—Dragon-flies.
(LIBELLULIDAE OF SOME AUTHORS) )
Elongate Insects with very mobile head and large eyes, with small
and inconspicuous antennae ending in a bristle; with four
elongate wings sub-equal in size and similar in texture, of
papyraceous consistency and having many veinlets, so that
there exists a large number of small cells. All the legs placed
more anteriorly than the wings. The earlier- stages of the
life are aquatic ; there is great change in the appearance of
the individual at the final ecdysis, but there is no pupal
enstar.
THE dragon-flies form a very natural and distinct group of
Insects. All the species are recognised with ease as belonging
to the family. They are invariably provided with wings in the
perfect state, and many of them are amongst the most active of
Insects. Their anatomy is, in several respects, very remarkable.
The head is large and is concave behind; it is attached to
the thorax in such a way that it rotates on two cervical sclerites
that project forwards, and in some cases almost meet in a point
in front; hence it possesses extreme mobility, the power of
rotation being very great.
The eyes are always large; in some cases they are even enor-
mous, and occupy the larger part of the area of the head: the
upper facets of the eye are in many cases larger than the
lower, and in a few forms the line of division is sharply marked
transversely. There are three ocelli, which, when the size of
the compound eyes is not too great, are placed in the usual
410 ; NEUROPTERA CHAP.
manner as a triangle on the vertex; but in the forms where
the compound eyes are very large the portion of the head
between is, as it were, puffed out so as to form a projection just
in front of where the eyes meet, and one ocellus is then placed
on each side of this projection, an antenna being inserted quite
close to it; the third ocellus is placed in front of the projection
Kae 7
0 Os SND fe oY
ap
a vl
a =
a)
Soy est Oke on,
SBE
=4|\— ase - —S
}
|
al : =
ad |
—_—
—— =
taal \ cael
Fic. 260.—Anaz formosus, Britain. (After Migneaux.) (The legs are not in a
natural position. )
we have mentioned, by which it is often much concealed; this
anterior ocellus is in some cases of unusually large size, and oval
or transverse in form.
The parts of the mouth are very peculiar, especially the
lower lip: we will briefly allude to its characters in the highly
modified forms, premising that in the smaller and less active
species it is less remarkable. The Libellulidae are carnivorous,
their prey being living Insects which are captured by the dragon-
fly on the wing; it is believed that the mouth is largely instru-
mental in the capture, though the flight of these Insects is so
excessively rapid that it is difficult, if not impossible, to verify
XVIII DRAGON-FLIES 411
the action of the mouthpieces by actual observation. For the
purpose of securing the prey a mouth that can change its
capacity to a considerable extent and with rapidity is a desider-
atum, and these qualities are present in thee mouths of those
Libellulidae that capture their prey while hawking. The upper lip
is very mobile, is pendent, and closes the mouth above, while the
lower lip entirely closes the under part by means of two mobile
plates; these in some forms (Libellula) meet together in the
mesial line, while in others a third plate separates them in the
middle (Fig. 261, B,/). These plates are, according to Gerstaecker’s
view, portions of the much changed labial palpi, the part that
separates them in Aeschna being the inner lobes of the labial max-
illae; in Libellula, where the dilated and valve-like joints of the
Fic. 261.—A, Maxilla
of Libellula quad-
rvimaculata ; B, la-
bium of Aeschna
grandis. Dp, Pp’,
Palpus; a, ter-
minal spine of
palpus ; ¢, cardo ;
t,stipes ; s,squama ;
le, outer lobe of
maxilla, partly
covered by, Ji,
inner lobe; 1m,
mentum ; 7, inter-
vening lobe. (After
Gerstaecker. )
palpi meet in the middle line, the labial lobes remain small and are —
overlapped by the dilated portions of the palpi. The maxillae
proper (Fig. 261, A) are less peculiar, their chief character being
that the inner and outer lobes are not separated, and that the palpus
is of only one joint. Some entomologists take, however, another
view of this structure, looking on the palp-like outer part (p of
our figure) as the true outer lobe of the maxillae, the palpus
proper being in that case considered to be entirely absent. The
mandibles are very powerful, and armed with largely developed
teeth. In the interior of the mouth there is a large, free, semi-
membranous lingua, the posterior part of its delicate inferior
lamina being connected with the mentum; the upper lamina of
the lingua is stronger and is pilose. The antennae of the dragon-
flies are always small, and consist of two stouter joints at the
1 Festschrift Ges. naturf. Freunde Berlin, 1873.
412 . NEUROPTERA CHAP.
base, and a terminal part which is very slender and pointed, and
formed of four or five joints.
The prothorax is always small; the pronotum is distinct,
though in some forms it is quite concealed in the concavity of
the back of the head; the sternum is small; the anatomy of the
pleura and basal pieces of the legs is obscure.
The meso- and meta-thorax are very intimately combined, —
and their relations are such that the former is placed much
above the latter. This.
peculiarity is carried to its
greatest extent in some of
the Agrioninae (Fig. 262,
A), where not only are
the wings placed at a
considerable distance be-
hind the three pairs of ©
legs, but also the front
pair of wings is’ placed
almost directly above the
hind pair. In the Anisop-
terides these peculiarities
are much less marked
(Fig. 262, B), nevertheless
even in them the three
pairs of legs are placed
quite in front of the wings.
This peculiar structure of
the wing-bearing segments
Fic. 262.—A, Agrion pulchellum, natural size ; B, 1s accompanied by an
Aeschna cyanea, profile ; C, same from front to unusual development of |
show position of legs. 4 natural size.
the pleura, which, indeed,
actually form the larger part, if not nearly the whole, of the
front region of the dorsal aspect of these two segments. We
shall not enter into more minute particulars as to the struc-
ture of the thorax, for difference of opimion prevails as to the
interpretation of the parts.’ The abdomen is remarkable for
its elongation ; it is never broad, and in some genera—AMecisto-
gaster, e.g.—it attains a length and slenderness which are not
1 Reference may be made to Calvert’s recent paper introductory to the study of
Odonata, in Jr. Amer. ent. Soc. xx. 1893, pp. 159-161.
XVIII DRAGON-FLIES 413
reached by any other Insects. It consists of ten segments and
a pair of terminal calliper-like or flap-like processes of very
various sizes and forms. |
The wings of the dragon-flies are usually transparent and
provided with a multitude of small meshes. The hind wings
are about as large as the front pair, or even a little larger; the
main nervures have a sub-parallel course, and are placed in
greater part on the anterior region of each wing. The relations
of the more constant nervures and the cells of which they are
parts form a complex subject, and are amongst the most im-
portant of the characters used in classifying these Insects. The
wings are always elongate in comparison to their’ breadth and
have no folds; they are held partially extended, or are placed
so as to project backwards, or backwards and outwards. They
exhibit another peculiarity, inasmuch as the front or costal
margin is slightly uneven before or near the middle, giving
rise to an appearance such as might result from the breaking
and subsequent mending of the marginal rib at the spot in
question, which is called the nodus. In some forms a peculiar
character exists in the shape of a small opaque space called the
membranule, lying close to the body of the Insect in the anal
area of the wing, as shown in Fig. 260.
The legs are slender and are chiefly remarkable for the
beautiful series of hair-like spines with which they are armed,
and which in some forms (eg. Platyenemis, Fig. 264) are of
considerable length. We believe that the legs are of great
importance in capturing the prey, they being held somewhat
in the position shown in Fig. 262, C. The tarsi are three-
jointed. In the male of Libellago caligata the legs exhibit a
remarkable condition, the tibiae being dilated, and on the upper
side of a vivid red colour, while below they are white. This
coloration and form are each unusual in the family. The male
of Platycnemis pennipes, a British species (Fig. 264), shows a
similar dilatation of the tibiae, but to a less extent and without
any great difference in the colour of the two faces of the dilata-—
tion. This dilatation reaches its maximum in Psiloenemis
dilatipes M‘Lach. ‘The position of the legs in relation to
the other parts of the body is peculiar to the dragon-flies; the
legs seem to be unfit for walking, the Insects never using them
for that purpose.
A414 NEUROPTERA CHAP,
Several peculiarities in the internal anatomy deserve notice.
The alimentary canal in Libellula is about as long as the body,
the oesophagus and chylific stomach being elongate, while the
intestine is short and divided into only two parts; there is no
definite proventriculus. The Malpighian tubules are shorter
than usual; they are about forty in number. The male has no
vesiculae seminales; the vasa deferentia are elongate, and the ©
ejaculatory duct is very short, being in fact merely a common
sinus formed by the terminations of the vasa deferentia. The
opening of this duct is situated on the penultimate ventral plate ;
the organs of intromission are, however, placed much anterior
to this, on the under side of the second segment. The mode in
which the fertilising fluid is transferred from the ninth to the
second segment is not well understood, but it is known that’
the abdomen is flexed by the Insect so as to bring the ninth
ventral plate into contact with the second. The three thoracic —
ganglia of the nervous chain are all contiguons, though not
completely amalgamated; the abdominal ganglia are seven in
number, and are all separated, the terminal one being larger than
the others. Dufour, after repeated dissections, was unable to find —
any salivary glands, but Olga Poletajewa' states that they exist.
The Odonata must. be ranked among the most highly-
organised Insects so far as external structure and powers of
locomotion are concerned; the peculiar modifications of the
thoracic segments and the relative positions of the wings and
legs mark a great departure from the normal type of Insect
structure. Their prey consists of living Insects, which they cap-
ture on the wing by their own superior powers of flight. They
destroy a great many Insects, their appetite for food being, as in
the cases of the Mantidae and of the tiger-beetles, apparently
almost insatiable. They are admirably constructed for the pur-
poses of their predatory lives; they fly with great swiftness and
change the direction of their flight with admirable facility.
They are, however, dependent on sunshine, and conceal them-
selves in dull and cloudy weather. The larger Insects of the
family belong to the division Anisopterides (Fig. 260, Anaxz
formosus), and some of these may, in our own country, usually be
seen, in the bright sunshine of the summer and autumn, engaged
in hawking in their favourite haunts. Places where other Insects
"1 Horae Soc. ent. Ross. xvi. 1881, p. 3.
“toy
XVII DRAGON-FLIES AI5
abound are naturally those most frequented; the glades of woods,
country lanes and hedge-sides, the borders of streams and the
margins of sheets of water are the places they most affect. They
inspire the rustics with some feeling of fear, and hence have
‘received the name of “ horse-stingers,’ and in North America are
called “devil's darning-needles.”. The aversion to dragon-flies
may perhaps be due to their appearance, which is certainly, in
the case of some of our species of Aeschna, Cordulegaster, and
Gomphus, very remarkable, consisting of a dark ground-colour
with bars and spots of vivid green or yellow, giving, it must be
admitted, a peculiar, even savage appearance to the Insects.
Whatever the reason may be, they are, it is certain, held in much
fear, and it is difficult to induce a country lad to touch one even
when it is captured and held by another person. The idea of
dragon-flies being dangerous to anything but their Insect victims
is, however, entirely erroneous; they may be captured and
handled without their inflicting any injury. It is probable that
the life of the imago may endure for several weeks if not months.
It is known that Sympyena fusca—a common European though
not British dragon-fly—hibernates in the imago state.
In the case of the large dragon-flies we have mentioned, each
‘ individual appears to have a domain, as it were, of its own.
Westwood tells us that he has seen what he believed to be the
same individual hawking daily for several weeks together over a
small pond. The writer observed a specimen of Cordulegaster
annulatus to frequent a particular bush, to which it returned
—frequently to the same leaf—after an excursion in search
of food. The way in which these Insects actually seize their
prey has not yet been made clear; it is certain that they
capture flying Insects, and it seems most probable, as we
have already said, that this is done by means of the legs.
These, as we have said, are inserted so as to be very near
to the mouth; they are directed forwards, and are held bent
at right angles so as to form a sort of net, and are armed
with a beautiful system of fine spines; it is probable that
if the dragon-fly pursue an Insect on the wing and strike it
with the trap, formed by its six.legs (Fig. 262, C), then these
immediately come together under the mouth, so that the victim,
directly it is captured by the leg-trap of its pursuer, finds
itself in the jaws of its destroyer. It is perhaps impossible to
416 NEUROPTERA CHAP,
verify this by actual observation, as the act of capture and trans-
fer is so very brief and is performed in the midst of a rapid
dash of flight, but it seems more probable that the prey is first
struck by the legs than that the mouth is the primary instrument
of capture. The excessive mobility of the head permits the victim
to be instantly secured by the mouth, and the captured fly is
turned about by this and the front pair of legs, and is nipped
rapidly so that the wings and drier parts fall off; the more
juicy parts of the prey are speedily squeezed into a little ball,
which is then swallowed, or perhaps we should rather say that
the mouth closes on it, and submits it to further pressure for the
extraction of the juices. We
have already noted that many of
these large and active dragon-
flies, particularly in the Libellu-
linae and Aeschninae, have their
eyes distinctly divided into two
parts, the facets in the lower
part of the eye being different
Exner considers * that the upper
movement, the lower for the
perception of the form of rest-
ing objects. Plateau thinks”
that the dragon-flies perceive
only movement, not form.
Fig. 263.—Inner view of a portion of . .
the left side of body of Libellula de- The splendid acts of flight
» press, showing a part of the mechanism of the Anisopterid Odonata are
of flight, viz. some of the chitinous . :
ridges at base of the upper wing, and accomplished by the aid of. a
some of the insertions of the tendons complex arrangement of chitin-
of muscles. A, line of section through :
base of upper wing, the wing being ous pleces at the bases of the
supposed to be directed backwards ; C, wings (Fig. 263). In Insects
upper portion of mechanism of the , :
lower wing ; 5, lever extending between with considerable powers of
the pieces connected with the two wings. 4; : Shae
(After von Lendenfeld, ) 8 flight the hind wings are usually
subordinate in functional im-
portance to the anterior, to which they are attached by a series
of hooks, or some other simple mechanism, on the wings.
1 Physiol. facett. Aug. 1891, p. 115.
2 Bull. Ac. Belgique (8), xvi. 1888, No. 11, p. 31.
from those of the upper part.
division is for the perception of -
XVIII DRAGON-FLIES 417
In the Odonata the two wings of each pair are quite free, but
they are perhaps brought. into correlative action by means of a
lever of unusual length existing amongst the chitinous pieces in
the body wall at the base of the wings (Fig. 263, >). The wing
muscles are large; according to von Lendenfeld* there are three
elevator, five depressor, and one adductor muscles to each wing:
he describes the wing movements as the results of the correlative
action of numerous muscles and ligaments, and of a great num-
ber of chitinous pieces connected in a jointed manner.
Amans?” has suggested that the mechanism of flight of the
dragon-fly would form a suitable model for a flying-machine, to
be propelled by electricity.
7 ED =
SS SNe
ee iw
SELES | \ SSS
CEES | 3 SS
Fic. 264.—Platycnemis pennipes, $, Britain.
The Zygopterides—the second of the two divisions of the’
Odonata—are Insects different in many respects from the large
and robust Anisopterides. The division comprises the delicate
Insects called “demoiselles,’ damsel-flies, by the French (Fig.
262, A,and Fig. 264). Great power of flight is not possessed by
these more fragile Insects; they flit about in the most gentle
and airy manner from stem to stem of the aquatic plants and
grasses that flourish in the localities they love. To this group
belong the fairy-like Insects of the genus Calepteryx, in which
various parts of the body and wings are suffused with exquisite
1 SB. Ak. Wien, \xxxiii. 1881, pp. 289-376, pls. i.-vii.
2 Rev. Sct, Nat. Montpellier (3), ii. p. 470.
VOL. V ) : ae Re
418 NEUROPTERA , CHAP.
metallic tints, while sometimes the two sexes of one species have
differently coloured wings. The smallest and most delicate
dragon-flies that are known are found in the tropics; some of
the genera allied to Agrion consist of Insects of extraordinary
fragility and delicacy.
Although the mature Odonata are so pre-eminently endowed
for an aerial and active life, yet in the earlier stages of their
existence they are very different; they are then, without excep-
tion, of aquatic habits; though carnivorous also in this period
of their existence, they are sluggish in movement, lurking in
concealment and capturing their prey by means of a peculiar
conformation of the mouth, that we shall subsequently describe.
Their life-histories are only very imperfectly known.
The éggs are deposited either in the water or in the stem of
some aquatic plant, the female Insect occasionally undergoing
submersion in order to accomplish the act. The young on
hatching are destitute of any traces of wings (Fig. 265), and the
structure of the thoracic segments is totally different from what
it is in the adult, the rectal respiratory system (Fig. 265, x), to
which we shall subsequently allude, being, however, already present.
The wings are said to make their first’ appearance only at the third
or fourth moult. At this time the pleura of the second and third
thoracic segments have grown ina peculiar manner so as to form
a lateral plate (Fig. 266, B, shows this plate at a later stage), and
the wing-pads appear as small projections from the membranes at
the upper margins of these pleural plates (Fig. 266, A, B). The
plates increase in size during the subsequent stadia, and meet
over the bases of the wing-pads, which also become much longer
than they were at first. The number of moults that occur during
growth has not been observed in the case of any species, but they
are believed to be numerous, There is no pupa, nor is there any
well-marked quiescent stage preceding the assumption of the
winged form at the last ecdysis, although at the latter part of its
life the nymph appears to be more inactive than usual. When full
grown, the nymph is more like the future perfect Insect than it was
at first, and presents the appearance shown in Figs. 266 and 270.
At this stage it crawls out of the water and clings to some sup-
port such as the stem or leaf of an aquatic plant; a few minutes
after doing so the skin of the back of the thoracic region splits,
and the imago emerges from the nymphal skin. The nymphs
a
XVIIr DRAGON-FLIES 419
never have the body so elongate as the perfect Insect, the differ-
ence in this respect being frequently great, and the nymphs of »
the subfamily Libellulinae being very broad (Fig. 266, nymph of
Ictinus sp.); consequently the creature on emergence from the
nymph-skin is very much shorter than it will soon become.
Fic. 265.—Larva of Diplax justhatched. n, Fie. 266.—Ictinus sp., nymph, Hima-
a ganglion of the ventral chain ; d, dorsal laya. is needless to
say that the instincts and stimuli connected with these migrations
are not understood. .
The nymphs are capable, under certain circumstances, of
accommodating themselves to very peculiar conditions of life.
The Sandwich Islands are extremely poor in stagnant waters, and
1 Rev. d@ Entomol. v. 1886, p. 232. * Riveau, Fewille Nat. xii. 1882, p. 123.
426 NEUROPTERA CHAP.
yet there exist in this remote archipelago several highly
peculiar species of Agrioninae. Mr. R. C. L. Perkins has
recently discovered that the nymphs of some
of these are capable of maintaining their
existence and completing their development
in the small collections of water that accumu-
late in the leaves of some lilies growing on
dry land. These nymphs (Fig. 271) have a
shorter mask than occurs, we believe, in
any other Odonata, and one would suppose
that they must frequently wait long for a
meal, as they must be dependent on stray
Insects becoming immersed in these tiny
reservoirs. The cannibal habits of the
te dha. Speen Odonata probably stand these lily-dwellers
short mask, living in in good stead; Mr. Perkins found that there
“ced ag ae were sometimes two or three nymphs of
different sizes together, and we may suspect
that it sometimes goes hard with the smaller fry. The extension
in the length of the body of one of these lily-frequenting Agrions
when it leaves the water for its aerial existence is truly extra-
ordinary.
The Odonata have no close relations with any other group of
Insects. They were associated by Latreille with the Ephemeridae,
in a family called Subulicornia. The members of the two groups
have, in fact, a certain resemblance in some of the features of
their lives, especially in the sudden change, without intermediate .
condition, from aquatic to aerial life; but in all important points
of structure, and in their dispositions, dragon-flies and may-flies
are totally dissimilar, and there is no intermediate group to
connect them. We have already said that the Odonata consist:
of two very distinct divisions—Anisopterides and Zygopterides.
The former group comprises the subfamilies Gomphinae, Cordu-
legasterinae, Aeschninae, Corduliinae, and Libellulinae,—Insects
having the hinder wings slightly larger than the anterior pair ;
while the Zygopterides consist of only two subfamilies—Calep- .
teryginae and Agrioninae; they have the wings of the two pairs
equal in size, or the hinder a little the smaller. The two groups
Gomphinae and Calepteryginae are each, in several respects, of
lower development than the others, and authorities are divided
a
XVIII DRAGON-FLIES 427
in opinion as to which of the two should be considered the more
primitive. It is therefore of much interest to find that there
exists an Insect that shares the characters of the two primitive
subfamilies in a striking manner. This Insect, Palaeophlebia
superstes (Fig. 272), has recently been discovered in Japan, and
‘is perhaps the most interesting dragon-fly yet obtained. De
Selys Longchamps refers it to the subfamily Calepteryginae, on
account of the nature of its wings; were the Insect, however,
deprived of these organs, no one would think of referring Palaeo-
phlebia to the group in question, for it has the form, colour, and
appearance of a Gomphine Odonate. Moreover, the two sexes
Fie. 272. — Palaeo-
phiebia superstes.
A, The Insect with
wings of one side
and with two legs
removed ; B, front
view of head of
female ; C,of male.
(After De Selys.)
ius
nui
differ in an important character—the form of the head and eyes.
In this respect the female resembles a Gomphine of inferior
development; while the male, by the shape and large size of
the ocular organs, may be considered to combine the characters
of Gomphinae and Calepteryginae. The Insect is very remark-
able in colour, the large eyes being red in the dead examples.
We do not, however, know what may be their colour during life,
as only one pair of the species is known, and there is no record
as to the life-history and habits. De Selys considers the nearest
ally of this Insect to be Heterophlebia dislocata, a fossil dragon-
fly found in the Lower Lias of England.
Numerous fossil dragon-flies are known; the group is well
represented in the Tertiary strata, and specimens have been
found in amber. In strata. of the Secondary age these Insects
weg iye ue NEUROPTERA CHAP. XVIII _
have been found as far back as the Lower Lias; their remains
are said to exist in considerable variety in the strata of that
epoch, and some of them to testify to the existence at that period
of dragon-flies as highly specialised as those now living. Accord-
ing to Hagen’ Platephemera antiqua and Gerephemera simplex,
two Devonian fossils, may be considered as dragon-flies; the
_ evidence as to this appears inadequate, and Brongniart refers the ~
latter Insect to the family Platypterides, and considers Plate-
phemera to be more allied to the may-flies.
One of the most remarkable of the numerous discoveries lately
made in fossil entomology is the finding of remains of huge Insects,
evidently allied to dragon-flies, in the Carboniferous strata at Com-
mentry. Brongniart calls these Insects Protodonates,’? and looks on
them as the precursors of our Odonata. Meganeura monyi was
the largest of these Insects, and measured over two feet across the
expanded wings. If M. Brongniart be correct in his restoration
of this giant of the Insect world, it much resembled our existing
dragon-flies, but had a simple structure of the thoracic segments,
and a simpler system of wing-nervures. On p. 276 we figured ©
Titanophasma fayoli, considered by Scudder and Brongniart as
allied to the family Phasmidae, and we pointed out that this
supposed alliance must at best have been very remote. This view
is now taken by M. Brongniart himself,? he having removed the
Insect from the Protophasmides to locate it in the Protodonates
near Meganeura. There appears to be some doubt whether the
wings supposed to belong to this specimen were really such, or
belonged rather to some other species.
1 Bull. Mus. Harvard, viii. 1880-81, p. 276.
* Insectes fossiles, p. 394. % Insectes fossiles, p..396.
CHAPTER XIX
AMPHIBIOUS NEUROPTERA CONTINUED——EPHEMERIDAE, MAY-FLIES
Fam. VII. Ephemeridae—May-flies.
Delicate Insects with atrophied mouth and small, short antennae ;
with four membranous wings
having much minute cross-
veining ; the hinder pair very
much smaller than the other
parr, sometimes entirely absent:
the body terminated by three
or two very elongate slender
tails. The earlier stages are
passed through in water, and
the individual then differs
greatly in appearance from
the winged Insect ; the passage
between the two forms is sud-
den; the creature in vts first
winged state is a subimago,
which by shedding a delicate
skin reveals the final form of
the individual.
THE may-flies are well known—in
literature—as the types of a_ brief
and ineffective life. This supposed
brevity relates solely to their existence in the winged form. In
the earlier stages the may-fly is so unlike its subsequent self
that it is not recognised as a may-fly by the uninitiated. The
total life of the individual is really quite as long as that of most
Fic. 273.—Ephemera danica, male,
Britain.
430 NEUROPTERA . CHa
other Insects. The earlier stages and life-histories of these
Insects are of great importance. The perfect Insects are so
delicate and fragile that they shrivel much in drying, and are
very difficult to preserve in a condition suitable for study.
The mouth of the imago is atrophied, the trophi scarcely
existing as separate parts. Packard says that in Palingenia
bilineata he could discover no certain traces of any of the mouth-
parts, but in Leptophlebia cupida he found, as he thought, the
rudiments of the maxillae and labium, though not of the mandibles.
The antennae are always short, and consist of one or two thick
basal joints sueceeded by a delicate needle-like segment, which,
though comparatively long, is
not divided. The ocular organs
are remarkable for their large
size and complex development ;
they are always larger in the
male than they are in the
female. The compound eyes of
the former sex are in certain
we species, e.g. Cloéon (Fig. 274),
Fia. 274.—Front of head of Cloéon, male. a, quite divided, so that each eye
Pillared eye ; 0, sessile eye ; ¢, ocellus. ‘
becomes a pair of organs of a
different character; one part forms a pillar facetted at its summit,
while the other part remains as a true eye placed on the side of
the head; in front of these compound eyes there are three ocelli.
Thus the Insect comes to have three different kinds of eyes,
together seven in number.
The prothorax is small, the pronotum being, however, quite
distinct. The mesothorax is very large; its notum forms by far
the larger part of the upper surface of the thoracic region, the
metathorax being small and different in structure, resembling
in appearance a part of the abdomen, so that the hind wings
look as if they were attached to a first abdominal segment. The
mesosternum is also disproportionately large in comparison with
the homologous piece preceding it, and with that following it.
The pleural ‘pieces are large, but their structure and disposition are
only very imperfectly understood. The coxae are small and are
widely separated, the anterior being, however, more elongate and
approximate than the others. The other parts of the legs are
slender ; the number of joints in the tarsi varies from five to one.
XIX . MAY-FLIES 431
The legs throughout the family exhibit a considerable variety of
structure, and the front pair in the males of some species are remark-
ably long. The abdomen is usually slender, and consists of ten
segments ; the terminal one bears three, or two, very long flexible
appendages. The first dorsal plate of the abdomen is either
wanting or is concealed to a considerable extent by the meta-
notum. The wings
are peculiar; the an-
terlor pair vary a
great deal in their
width, but are never
very long in propor-
tion to the width;
the hind pair are
always dlispropor-
tionately small, and
sometimes are quite
wanting. The vena-
tion consists of a few, or of a moderate number, of delicate longi-
tudinal veins that do not pursue a tortuous course, but frequently
Fig. 275.—Wings of Ephemera danica. (After Eaton.)
are gracefully curved, and form a system of approximately similar
curves, most of the veins being of considerable length ; close to the
anterior margin of the wing there are two or three sub-parallel
veins. Frequently there are very numerous fine, short cross-
veinlets, but these vary greatly and may be entirely wanting.
The earlier stages of the life of Ephemeridae are, it is believed,
in the case of all the species, aquatic. May-flies, indeed, during
the period of their post-embryonic development are more modified
for an aquatic life than any other Insects, and are provided with
a complex apparatus of tracheal gills. The eggs are committed
to the waters without any care or foresight on the part of the
parent flies, thus the embryonic development is also aquatic ;
little, however, is known of-it. According to Joly* the process
in Palingenia virgo is slow. The larva on emerging from the
egg has no respiratory system, neither could Joly detect any
circulation or any nervous system. The creature on emergence
is very like Campodea in form, possessing long antennae and tails
—caudal setae. Owing to the organisation being inferior, the
creature in its earlier stages is called a larvule; in its later stages
1 Mem. Ac. Sci. Toulouse (7), iii. 1871, p. 379.
432 NEUROPTERA CHAP.
it is usually spoken of as a nymph, but the term larva is also
frequently applied to it. Soon the gills begin to appear in the
form of small tubular caeca placed in the posterior and upper
angles of the abdominal rings; in fifteen days the gills begin to
assume their characteristic form, are penetrated by tracheae, and
e
\
¥
SOSS esa
~eee o
oF
GT 4 MAE
é ka
Fia.276.—Nymph of Cloéon dipterum.' Wing-sheath Fic. 277.—Larvule of Oloéon
of left side, gills of right side, removed ; g, dimidiatum. (After Lub-
tracheal gills. (After Vayssiére.) bock.)
the circulation can be seen. The amount of growth accomplished
after hatching between March and September is but small.
The metamorphosis of Cloéon has been described by Sir John
Lubbock; he informs us that the young creature undergoes a
constant and progressive development, going through a series of
more than twenty moults, each accompanied by a slight change
of form or structure. His observations were made on captured
' In reference to a doubt as to the name of this nymph cf. Eaton, 77. Linn.
Soc. Zool, (2) ili, p. 20.
ia.’
13
tN ga SD
XIX MAY-FLIES 433
specimens, so that it is not certain that what he calls' the first
stage is really such. He found no tracheae in the earliest stages ;
the small first rudiments of the gills became visible in the third
stage, when there were no tracheae; the fourth instar possessed |
tracheae, and they could be seen in the gills. The wing rudi-
ments could first be detected in the ninth and tenth stages. The
changes of skin during the winter months are separated by
longer intervals than those occurring at other periods of the year.
The nymphs differ greatly in the structure and arrangement
of their tracheal gills, and display much variety in their general
form and habits; some of them are very
curious creatures. Pictet? divides them
in accordance with their habits into four
groups: (1) Fossorial larvae: these live
in the banks of streams and excavate
burrows for ‘shelter; they are of cylin-
drical form, possess robust legs, abundant
gills at the sides of the body, and
frequently processes projecting forwards
from the head: examples, Lphemera (Fig.
278) and Palingenia. (2) Flat larvae:
these live attached to rocks, but run with
rapidity when disturbed ; they prefer rapid
streams, have the breathing organs at-
tached to the sides. of the body and not
reposing on the back; they are exclu-
sively carnivorous, while the fossorial
forms are believed to obtain their nutri-
ment by eating mud: example, Baétis.
(3) Swimming larvae: elongate delicate
creatures, with feeble legs, and with strongly |
ciliated caudal setae: example, Cloéon (Fig.
276). (4) Climbing larvae: these live in
slowly-moving waters, especially such as
have much slimy mud in suspension, and yy, 278,—Adult nymph of
they have a habit of covering them- welts vulgata. (After
2 : , aton.) Britain.
selves with this mud sometimes to such
an extent as to become concealed by it: example, Potamanthus.
1 Tr. Linn. Soc. xxiv. 1863, p. 62, and xxv. 1866, p. 477.
-2 Hist. Nat, Newropt. Ephémérines, 1843, p. 24.
bo
os
VOL. V
434 me NEUROPTERA CHAP.
~The anatomy of the nymphs has been treated by Vayssiére,"
who arranges them in five groups in
accordance with the conditions of the
tracheal gills: (1) The gills are of large
size, are exposed and furnished at the
sides with respiratory fringes: ex-
ample, Ephemera (Fig. 278). (2) The
branchiae are blade-like, not fringed,
and are exposed at the sides of the
body: example, Cloéon (Fig. 276). (3)
The respiratory tubes are placed on
the under surface of plates whose —
upper surface is not respiratory: ex-
ae eal, ample, Oligoneuria garumnica (Fig.
xt 279). (4) The anterior gill is modi-
a: fied to form a plate that covers the
others: example, Tricorythus (Fig.
282,B). (5) The gills are concealed in
a respiratory chamber : example, Proso-
pistoma (Fig. 280). The last of these
Fin. Ste /aeieveneh. Gt Oligoneuria nymphs is more completely adapted
garumnica, France. gz and g,, for an aquatic life than any other
tee dorsal tracheal gills. Tnsect: at present known; it was for
yssiere. )
long supposed to be a Crustacean, but
it has now been shown to be the early stage of a may-fly,
the sub-imago having been reared from the nymph. The
carapace by which the larger part of the body is covered is
formed by the union of the pro- and meso-thorax with the sheaths
of the anterior wings, which have an unusually extensive develop-
ment; under the carapace there is a respiratory chamber, the
floor and sides of which are formed by the posterior wing-
sheaths, and by a large plate composed of the united nota of the
metathorax and the first six abdominal segments. In this
chamber there are placed five pairs of tracheal gills; entrance of
water to the.chamber is effected by two laterally-placed orifices,
and exit by a single dorsal aperture. These nymphs use the
body as a sucker, and so adhere strongly to stones under water.
When detached they swim rapidly by means of their caudal
setae; the form of these latter organs is different from that
1 Ann. Sci. Nat. Zool. (6) xiii. 1882, pp. 1-137, pls. 2-11.
XIX MAY-FLIES 435
of other Ephemerid nymphs. This point and other details
of the anatomy of this creature have been
described -in detail by Vayssiére.’ These
nymphs have a very highly developed tracheal
system; they live in rapid watercourses
attached to stones at a depth of three to six
inches or more under the. water. Species of
Prosopistoma oceur in Europe, Madagascar,
and West Africa.
. According to Eaton, in the nymphs of
some Ephemeridae the rectum serves, to a
certain extent, as a respiratory agent ; he con-
siders that water is admitted to it and ex-
pelled after the manner we have described in a, yh i
Odonata, P: £21. , Fia. 280. — Prosopi-
The internal anatomy of the nymphs cf stoma punetifrons,
Ephemeridae shows some points of extreme Y™Ph. France.
; (After Vayssiére.) 0,
interest. The long Orifice of exit from
+ :
Ll eben Y caudal setae are — T@spitatory chamber.
respiratory organs of a kind that
a ae is almost if not quite without
parallel in the other divisions of
+ SN i Insecta. The dorsal vessel for the
circulation of the blood is elongate,
aa Hy rst and its chambers are arranged one
Fic. 281.—A, Last three abdominal to each segment of the body. It
segments and bases of the three drives the blood forwards in the usual
caudal processes of Cloéon dip- .
ferum: +, dorsal vessel : kl, ostia MANNer, but the posterior chamber
thereof ; %, special terminal cham- possesses three blood-vessels, one of
ber of the dorsal vessel with its , . :
entrance « ; b, blood-vessel of the Which is prolonged into each caudal
. . av agioats prone H B, Ai pe seta. This terminal chamber is so
= S1X olnt O e€ leit caudal pro- °
: ews are below ; b, a Cortina of arranged as to drive the blood back-
the blood-vessel ; 0, orifice in the wards into the vessels of the setae;
latter. (After Zimmermann.) ‘
: on the under surface of the vessels
there are oval orifices by which the blood escapes into the
cavity of the seta so as to be submitted to the action
of the surrounding medium for some of the purposes of
respiration. This structure has been described by Zimmer-
1 Ann. Sci. Nat. Zool. (7) ix. 1890, pp. 19-87, pls. 2-5.
2 Ann. Nat. Hist. (3) xviii. 1866, p. 145.
' Fic. 282.—A, Nymph of Ephemerella ignita
436 NEUROPTERA CHAP.
mann,’ who agrees with Creutzberg ’ that the organ by which the
blood is propelled into the setae is a terminal chamber of the
dorsal vessel ; Verlooren,* who first observed this accessory system
of circulation, thought the contractile chamber was quite separate
from the heart. The nature of the connexion between this
terminal chamber that drives the blood backwards and the
other chambers that propel the fluid forwards appears still to
want elucidation.
The nymphs of the Ephemeridae being creatures adapted for
existence in water, the details
of their transformation into
creatures having an entirely
aerial existence cannot but. be
of much interest. In the
nymphs the tracheal system is
well developed, but differs from
that of air-breathing Insects
in the total absence of any
spiracles. Palmén has inves-
tigated this subject,* and finds
that the main longitudinal
tracheal trunks of the body of
the nymph are not connected
with the skin of the body by
tracheae, but are attached
thereto by ten pairs of slender
strings extending between the
with gills of left side removed; g, gills: Chitinous integument and the
B, nymph of Tricorythus sp. with gill tracheal trunks. When the
ae aie Mey gill covers skin is shed these strings—or
- rather a chitinous axis in each
one—are drawn out of the body, and bring with them the chitinous.
A A 4 B
Sa:
linings of the tracheae. Thus notwithstanding the absence of spir-
acles, the body wall is at each moult pierced by openings that
extend to the tracheae. After the ordinary moults these orifices close
immediately, but at the change to the winged state they remain open
and form the spiracles. At the same time the tracheal gills are com-
1 Zeitschr. wiss. Zool. xxxiv. 1880, p. 404.
2 Ann. Nat. Hist. (5) xv. 1885, p. 494. 3 Mem. Cour. Ac. Belg. 4to, xix. 1847, p. 1.
4 Zur Morphologie des Tracheensystems, Helsingfors, 1877, pp. 1-20.
XIX MAY-FLIES 437
pletely shed, and the creature is thus transformed from a water-
breather to an Insect breathing air as usual. In addition to this
change there are others of great importance, such as the develop-
ment of the great eyes and the complete atrophy of the mouth-
parts. The precise manner of these changes is not known; they
occur, however, within the nymph skin. The sudden emergence
of the winged Insect from the nymph is one of the most
remarkable facts in the life-history of the may-fly; it has been
observed by Sir John Lubbock,’ who describes it as almost in-
stantaneous. The nymph floats on the water, the skin of the back
opens, and the winged Insect flies out, upwards and away ;
“from the moment when the skin first cracks not ten seconds are
over before the Insect has flown away.” The creature that thus
escapes has not, however, quite completed its transformation. It
is still enveloped in a skin that compresses and embarrasses it ;
this it therefore rapidly gets rid of, and thus becomes the
imago, or final instar of the life-cycle. The instar in which the
creature exists winged and active, though covered with a skin, is
called the sub-imago. The parts of the body in the sub-imago are
as a whole smaller than they are in the imago, and the colour is
more dingy; the appendages—wings, legs, and caudal setae—are
generally considerably shorter than they are in the imago, but
attain their full length during the process of extraction. The
creatures being, according to Riley, very impatient and eager to
take to the wing, the completion of the shedding of the skin of
the sub-i “imago is sometimes performed while the Insect is flying
in the air.
The food of young Rah auenie is apparently of a varied and
mixed nature. Eaton says”
that though sometimes the
stronger larvae devour the
weaker, yet the diet is even
in these cases partly. vege- Ss
table. The alimentary canal
frequently contains much
mud ; VESEY: small organisms, Fig. 283.—Lingua of Heptagenia longicauda,
such as diatoms and con- x16. m, Central; /, lateral pieces. (After
: Vayssiére.)
fervae, are thought to form
a large part of the bill of fare of Ephemerid nymphs. Although
1 Tr. Linn. Soc, xxv. 1866, p. 483. 2 Ann. Nat. Hist. (3) xviii. 1866, p. 145.
438 | NEUROPTERA CHAP.
the mouth is atrophied in the imago, yet it is highly
- developed in the nymphs. This is especially notable in the
case of the lingua or hypopharynx (Fig. 283); indeed Vayssiére *
seems to incline to the opinion that this part of the mouth may
be looked on in these Insects as a pair of appendages of a head-
segment (see p. 96 ante), like the labium or maxillae. |
The life-history has not been fully ascertained in the case of
any species of may-fly ; it is known, however, that the develop-
ment of the nymph sometimes occupies a considerable period, and
it is thought that in the case of some species this extends to
as much as three years. It is rare to find the post-embryonie
development of an Insect occupying so long a period, so that we
are justified in saying that brief as may be the life of the may-
fly itself, the period of preparation for it is longer than usual.
Réaumur says, speaking of the winged fly, that its life is so short
that some species never see the sun. Their emergence from the
nymph-skin taking place at sunset, the duties of the generation
have been, so far as these individuals are concerned, completed before
the morning, and they die before sunrise. He thinks, indeed,.
that individuals living thus long are to be looked on as Methuselahs
among their fellows, most of whom, he says, live only an hour or
half an hour. It is by no means clear to which species these
remarks of Réaumur refer; they are doubtless correct in certain
cases, but in others the life of the adult is not so very short, and
in some species may, in all probability, extend over three or four —
days; indeed, if the weather undergo an unfavourable change so
as to keep them motionless, the life of the flies may be prolonged
for a fortnight.
The life of the imago of the may-fly is as remarkable as it is
brief; in order to comprehend it we must refer to certain peculi-
arities of the anatomy with which the vital phenomena are con-
nected. The more important of these are the large eyes of the
males, the structure of the alimentary canal, and that of the
reproductive organs. We have already remarked that the parts
of the mouth in the imago are atrophied, yet the canal itself not
only exists but is even of greater capacity than usual; it appears
to have much the same general arrangement of parts as it had in
the nymph. Its coats are, however, of great tenuity, and according
1 Ann. Sci. Nat. Zool. (6) xiii. 1882, p. 118.
2 Réaumur, A/em. vi. 1742, p. 457.
XIX MAY-FLIES 439
to Palmén? the divisions of the canal are separated by changes
in the direction of certain portions anterior to, and of others
posterior to, its central and greater part—the stomach—in such
a manner that the portions with diverted positions act as valves.
The stomach, in fact, forms in the interior of the body a delicate
capacious sac; when movement tends to increase the capacity of
the body cavity then air enters into the stomachic sac by the
mouth orifice, but when muscular contractions result in pressure
on the sac they close the orifices of its extremities by the valve-
like structures we have mentioned above; the result is, that as
complex movements of the body are made the stomach becomes
more and more distended by air. It was known even to the old
naturalists that the dancing may-fly is a sort of balloon, but they
were not acquainted with the exact mode of inflation. Palmén
says that in addition to the valve-like arrangements we have
described, the entry to the canal is controlled by a circular muscle,
with which are connected radiating muscles attached to the walls
of the head. Palmén’s views are adopted, and to a certain extent
confirmed, by Fritze,? who has examined the.alimentary canal of
the may-fly, and considers that though the normal parts of the
canal exist, the function is changed in the imago, in which the
canal serves as a sort of balloon, and aids the function of the
reproductive organs. The change in the canal takes place in an
anticipatory manner during the nymph and sub-imago stages.
The sexual organs of Ephemeridae are remarkable for their
simplicity ; they are destitute of the accessory glands and diver-
ticula that, in some form or other, are present in most other
Insects. Still more remarkable is the fact that the ducts by
which they communicate with the exterior continue as a pair to the
extremity of the body, and do not, as in other Insects, unite into
a common duct. Thus in the female there is neither bursa copu-
latrix, receptaculum seminis, nor uterine portion of oviduct, and
there is no trace of an ovipositor; the terminations of the ducts
are placed at the hind margin of the seventh ventral plate, just
in front of which they are connected by a fold of the integu-
ment. The ovary consists of a very large number of small egg-
tubes seated on one side of a sac, which forms their calyx, and
one of whose extremities is continued backwards as one of the
1 Uber paarige Ausfiihrsginge, etc., Helsingfors, 1884, p. 53.
2 Ber. Ges. Freiburg, iv. p. 5; ef. J. R. Mier. Soc. 1889, p. 206.
440 NEUROPTERA CHAP.
pair of oviducts. The male has neither vesiculae seminales, acces-
sory glands, nor ductus ejaculatorius. The testes are elongate
sacs, whose extremities are prolonged backwards forming the vasa
deferentia; these open separately at the extremity of the body, —
each on a separate intromittent projection of more or less complex
character, the two organs being, however, connected by means of —
the ninth ventral plate, of which they are, according to Palmén,
appendages. We should remark that this authority considers
Heptagenia to form, to some extent, an exception as regards the
structures of the female; while Polymitarcys is in the male sex
strongly aberrant, as the two vasa deferentia, instead of being
approximately straight, are bent inwards at right angles near
their extremities so as to meet, and form in the middle a common
cavity, which then again becomes double to pass into the pair of —
intromittent organs.
According to the views of Exner and others, the compound
eyes of Insects are chiefly organs for the perception of movement ;
if this view be correct, movements such as those made during
the dances of may-flies may, by the number of the separate eyes, by
their curved surfaces and innumerable facets, be multiplied and
correlated in a manner of which our own sense of sight allows
us to form no conception. We can see on a summer's evening how
beautifully and gracefully a crowd of may-flies dance, and we may
well believe that to the marvellous ocular organs of the flies them-
selves (Fig. 274) these movements form a veritable ballet. We
have pointed out that by this dancing the peculiarly formed aliment-
ary canal becomes distended, and may now add that Palmén and
Fritze believe that the unique structure of the reproductive organs
is also correlated with the other anatomical peculiarities, the con-
tents of the sexual glands being driven along the simple and
direct ducts by the expansion of the balloon-like stomach. During
these dances the momentary conjugation of the sexes occurs,
and immediately thereafter the female, according to Eaton,
resorts to the waters appropriate for the deposition of her eggs.
As regards this, Eaton says:* “Some short-lived species discharge
the contents of their ovaries completely en masse, and the pair
of fusiform or subcylindrical egg-clusters laid upon the water
rapidly disintegrate, so as to let the eggs sink broadcast upon
the river-bed. The less perishable species extrude their eggs
1 Tr. Linn, Soc. 2nd ser. Zool. iii. 1883, p. 11.
SSE
XIX MAY-FLIES 441
eradually, part at a time, and deposit them in one or other of
the following manners: either the mother alights upon the
water at intervals to wash off the eggs that have issued from
the mouths of the oviducts during her flight, or else she creeps
down into the water to lay her eggs upon the under-side of
stones, disposing them in rounded patches, in a single layer
evenly spread, and in mutual contiguity.” The eggs are very
numerous, and it is thought may sometimes remain in the
water as much as six or seven months before they hatch.
The number of individuals produced by some kinds of may-
flies is remarkable. Swarms consisting of millions of individuals
are occasionally witnessed. D’Albertis observed Palingenia
papuana in countless myriads on the Fly River in New Guinea:
“For miles the surface of the river, from side to side, was white
with them as they hung over it on gauzy wings; at certain
moments, obeying some mysterious signal, they would rise in
the air, and then sink down anew like a fall of snow.” He
further states that the two sexes were in very disproportionate
numbers, and estimates that there was but a single female to
every five or six thousand males.
Ephemeridae in the perfect state are a favourite food of
fishes, and it is said that on some waters it is useless for the
fly-fisher to try any other lure when these flies are swarming.
Most of the “duns” and “spinners” of the aigler are
Ephemeridae ; so are several of the “ drakes,” our large ZL. danica
and #. vulgata being known as the green drake and the gray drake.
Ronalds says’ that the term “dun” refers to the pseud-imago
condition, “spinner” to the perfect Insect. . danica and E£.
vulgata are perhaps not distinguished by fishers;. Eaton says
that the former is abundant in rapid, cool streams, while JZ.
vulgata prefers warmer and more tranquil rivers.
These sensitive creatures are unable to resist the attractions
of artificial lights. Réaumur noticed this fact many years ago,
and since the introduction of the electric light, notes may
frequently be seen in journals recording that myriads of these
Insects have been lured by it to destruction. Their dances may
frequently be observed to take place in peculiar states of light
and shade, in twilight, or where the sinking sun has its light
rendered broken by bushes or trees; possibly the broken lights
1 Fly-Fisher’s Entomology, 4th ed. 1849, p. 49.
442 NEUROPTERA CHAP,
are enhanced in effect by the ocular structures of the Insects. |
It has recently been ascertained that a species of Zeleganodes
is itself luminous. Mr. Lewis,' who observed this Insect in
Ceylon, states that in life the whole of the abdomen was lumin-
ous, not brightly so, but sufficient to serve as a guide for captur-
ing the Insect on a dark night. It has also been recorded that
the male of Caenis dimidiata gives a faint blue light at night.
Nearly 300 species of Ephemeridae are known, but this
may be only a fragment of what
‘ foi actually exist, very little being
known of may-flies of other
parts of the world than Europe
and North America. One of the
more curious forms of the family
is Oniscigaster wakefieldi; the
body of the imago is unusually
rotund and furnished with lateral
processes. In Britain we have
about forty species of may-fly.
The family is treated as a distinct
Order by Brauer and Packard, and
is called Plectoptera by the latter.
That Insects so fragile, so
highly organised, with a host of
powerful enemies, but themselves
destitute of means of attack or
Fig, 284.—Oniscigaster wakefieldi. New defence, should contrive to exist
. Zealand. (After M‘Lachlan.)
nah
pe\\
Tr
oN
wf
Gina i
Ns
pun,
afi
appears still more unlikely that such delicate Insects as
Ephemeridae should leave implanted in the rocks their traces
in such a manner that they can be recognised; nevertheless,
such is the case,—indeed, the may-fly palaeontological record is
both rich and remarkable. Several forms are preserved in
amber. In the Tertiary bed of the old lake at Florissant, Scudder
has been able to distinguish the remains of no less than six
species; while in the Jurassic layers of ‘the Secondary epoch, in
more than one locality, the remains of several other species
have been detected and described. Still more remarkable is the
fact that in the Devonian and Carboniferous layers of the
1 P. ent. Soc. London, 1882, p. xiii.
at all is remarkable; and it >
*
2 a « - >
ee ~ =
> fe
Mies
5 EE er ee
XIX _ MAY-FLIES 443
Palaeozoic period, remains are found that appear to be akin to
our existing Ephemeridae. Palingenia feistmantelic from the
Carboniferous of Bohemia is actually referred to a still existing
genus; it is said to have been of gigantic size for a may-fly.
The families Megasecopterides, Platypterides, and Stenodicty-
opterides of the Carboniferous epoch (see p. 343) are all more or
less closely allied to the Ephemeridae, and in addition to these
Brongniart has established the family Protephemerides for some
Insects that he considers to have been the precursors in the
Carboniferous epoch of our existing
may-flies. These ancient Insects
differed in having the wings of
another form from those of exist-
ing Ephemeridae, and in having
the hind wings equal in size to
the front pair. Besides this, these
Insects had, as shown in Fig. 285,
prothoracic dorsal appendages ;
some had also projections from the
abdominal segments, considered by
Brongniart to be of the nature-of
gills. Some doubt must exist as to
this point, for we find in the imago
of one of our existing Ephemeridae,
Oniscigaster wakefieldi, Fig. 284,
abdominal processes that are not
gills.
It ‘is remarkable that may-
flies, which now form a com-
paratively unimportant part of
the Insect tribe, should in far
distant times have been represented Fic. 285.—Homaloneura bonnieri ; Car-
by so great a variety of allied forms. — boniferous of Commentry, (After
Our fragile, short-lived may-flies wien
appear to be, as Scudder says, the lingering fragments of an
expiring group.
CHAPTER XX
NEUROPTERA PLANIPENNIA——SIALIDAE, ALDER-FLIES, SNAKE-FLIES—_
PANORPIDAE, SCORPION-FLIES — HEMEROBIIDAE, ANT-LIONS,
LACEWINGS, ETC.
Fam. VIII. Sialidae—Alder-flies and Snake-flies.
Four wings of moderate size, meeting in repose over the back at
an angle; the hinder of the two pairs slightly the smaller ;
the anal area small or nearly absent, not plicate. Nervures
moderately numerous, transverse veinlets moderately numerous,
forming irregularly disposed cells. The metamorphosis is
great; there is a quiescent pupa. Thelarvahas the mandibles
JSormed for biting, armed with strong teeth. | |
THE Sialidae, though but a small family of only some six or
eight genera, comprise
a considerable variety of — -
forms and two sub- —
families — Sialides and
Raphidiides. The former
eroup has larvae with
aquatic habits possessed
of branchiae but no
spiracles.
Sialis lutaria is one
of the commoner British
Insects frequenting the
a vegetation about the
tn banks of tranquilstreams;
Pi 286 Mh ey, int din, Bei. ig well known 8
anglers, being used by
them for a bait. According to Ronalds it is called the alder or
é
nae ee ee ey ee
= 5
=e
CHAP. XX» SIALIDAE 445
orl-fly, and in Wales the humpback. It is very unattractive in
appearance, being of a _ blackish colour, with wings of a
yellow-brown tinge, and makes but a poor show when flying.
The female deposits patches of elongate eggs, placed on end and
packed together in a very clever manner (Fig. 287). These patches
of eggs, of a stone-gray colour, are common objects on rushes
or stems of grass near water, and it is stated that there may
be no less than 2000 or 3000 eggs in one of them. Our
figure gives some idea of the mode in which the eggs are arranged,
Fic. 287.—Portion of a row of eggs of Fig. 288.—Sialis lutaria,
Sialis lutaria. (After Evans.) larva.
and the curious narrow process that exists at the end of each.
The eggs are said to be sometimes placed at a considerable distance
from water, so that when the tiny larvae are hatched they
must begin their lives by finding the way to a suitable pool or
stream. The larvae (Fig. 288) are objects of very great interest
owing to each of segments 1 to 7 of the hind body- being furnished
on each side with a jointed filament, while the last segment ends
in a still longer, but unjointed process. These filaments are
branchiae by means of which the Insect obtains air, being, as we
have said, destitute of spiracles. It is an active creature and
waves its filaments in a very graceful manner; this process no
doubt aids the branchiae in their respiratory work. These larvae
are well able to exist out of water if they have a sufficiently
damp environment. They live on animal matter, but their life-
history has not been followed in much detail and it is not known
446 NEUROPTERA - CHAP.
how many moults they make. The young larva has the head.
disproportionately large and the branchial filaments longer.
When the growth is completed the larva returns to land, seeks a
suitable situation in the soil, and after an interval changes to a
pupa, in which the characters of the perfect Insect are plainly
visible. Subsequently, without becoming again active, it changes
to the perfect Insect, and enjoys, for a few days only, an aerial life.
The anatomy of the larva has been treated by Dufour.’ The
supra-oesophageal ganglion is remarkably small; nothing is. said
as to the existence of an infra-oesophageal ganglion; there are
three thoracic and eight abdominal
5 te ganglia; the first pair of these latter
: Te ff y. axe nearer together than the others, and
a A f= this is also the case with the last three.
PE The alimentary canal in the adult is
a provided with a large paunch attached
to the crop by a narrow neck, but
chiae has also been described by the
indefatigable French entomotomist. A
tracheal tube sends a branch into one
of. the appendages (Fig. 289);
branch gives off numerous smaller
tracheae, which at their extremities
break up into branchlets close to the
integument. The tracheal tube that
receives each main branchial trachea,
sends off from near the point of entry
Fic. 289.—Structure of tracheal gill Of the latter another trachea, that
: “56% the ae ae distributes its branchlets on the ali-
trunk with which it is con- mentary canal. The margins of each
by altvnetaey Oeeaee given off» ypendage are set with swimming hairs,
so that the branchiae act as organs of
locomotion as well as of respiration, and by their activity in the
former capacity increase the efficiency of their primary function.
The genus Sialis occurs in a few species only, throughout the
1 Ann. Sci. Nat. series 8, ix. Zool. 1848, p. 91, pl. 1.
? Newport, Zr. Linn. Soc. xx. 1851, pl. 21, fig. 13. Loew, however, who also
describes and figures the anatomy of s. lutaria, states that there is no paunch,
Linnaea entomologica, iii, 1848, p. 354.
Dufour could find no trace of this in °
the larva. The structure of the bran-—
pr, Oa
Reet, : | | SIALIDAE 447
whole of the Palaearctic and Nearctic regions, and reappears in
Chili, though absent in all the intervening area: Several other
. genera of Insects exhibit the same peculiarity of distribution.
The genera Corydalis and Chauliodes form a group distinct
from Stalis, and are totally differ- |
ent in appearance, being gigantic ——— ; i ie
Insects, sometimes with the man-
fi
dibles of the male enormously
elongated (Fig. 290). The species
of Corydalis are called in North
America Hellgrammites; Riley
has described and figured the
metamorphosis of C. cornutus, the
life-history being very similar to
that of our little Sialis. A mass
consisting of two or three thousand
eggs is formed by the female, and
the young larva has long fila-
ments at the sides of the body
like Sialis. These in the later
larval life are comparatively shorter,
but the Insect is then provided
with another set of gills in the Fre, 290.—Corydalis crassicornis, male,
form of spongy masses on the ith grater portions of te wing
under-side of the body. Riley, .
however, considers that these organs serve the purpose of attach-
ment rather than ‘of respiration. The
larvae are known to the Mississippi
fishermen as crawlers, and are greatly
esteemed as bait.
The Raphidiides. or snake-flies form
the second tribe of Sialidae. There are
only two genera, Raphidia and Inocellia,
| peculiar to the Palaearctic and Nearctic
Fig. 291.—Raphidia notata, fe- regions. The perfect Insects are chiefly
male. Britain, (After Curtis.) :
: remarkable for the elongation of the
prothorax and back of the head to form a long neck, and for
the existence in the female of an elongate exserted ovipositor.
1 M‘Lachlan, Ent. Month. Mag. vii. 1870, p, 145.
2 Rep. Ins. Missouri, ix. 1877, p. 125.
448 NEUROPTERA CHAP.
The species are rather numerous, and have been recently
monographed by Albarda.’ The three or four British species
of the genus are all rare Insects, and occur only in wooded
regions,
The Raphidiides, like the Sialides, have a carnivorous larva,
which, however, is terrestrial in habits, feeding, it would appear,
chiefly on Insects that harbour in old timber. The snake-fly
larvae (Fig. 292) are very ingenious in their manner of escaping,
which is done by an extremely rapid wriggling backwards. They
are capable of undergoing very prolonged
fasts, and then alter in form a good deal,
becoming shorter and more shrivelled;
Fig. 292 is taken from a specimen that
had been fasting for several weeks. They
are excessively voracious, and hunt after
the fashion of beasts of prey; their habits
have been described by Stein,? who states
that he kept a larva from August to the
end of May of the following year without
food ; it then died in a shrivelled-up state.
The larva of the snake-fly changes to a
pupa that is remarkably intermediate in
form between the perfect Insect and the
larva; the eyes, legs, wing-pads, and ovi-
positor being but little different from those
of the imago, while the general form is
Bro, 292.—Raphidia notaity that of the larva, and the peculiar elonga-
arva. New Forest. 2 “ .
tion of the neck of the imago is absent.
This pupa differs from that of Sialis in the important particular
that before undergoing its final ecdysis it regains its activity and
is able to run about.
The internal anatomy of Raphidia has been treated by Loew,*
and is of a very remarkable, character ; we can here only mention
that the salivary glands consist of a pair of extremely elongate
tubes, that there is a very definite paunch attached as an ap-
pendage to one side of the crop, and that the most peculiar
character consists of the fact that, according to Loew, four of the
six Malpighian tubes have not a free extremity, being attached
-
a
aM TN
giill|
|
Mb
Nit
as
i
=
Ar
Te
oO
+
Ls
Kua be
Aad eC
1 Tijdschr. Ent. vol. xxxiy. 1891. 2 Arch. f. Naturg. iv. i. 1838, p. 315.
> Linnaea entomologica, iii. p. 1848, 346, pl. i.
=
Y
XX SIALIDAE AND SCORPION-FLIES 449
at each end so as to form elongate loops; the mesenteron is very
complex in character.
A considerable number of fossil re-
mains from both Tertiary and Mesozoic .
strata are referred to Sialidae; and a
laryal form from the red_ sandstone
of Connecticut has been considered by
Seudder to be a Sialid, and named
Mormolucoides articulatus, but the cor-
rectness of this determination is - very
doubtful (Fig. 293). These fossils are,
however, of special interest as being the
most ancient Insect larvae yet brought .
to light. A still older fossil, from the Car-
boniferous strata of Illinois called Miamia
bronsont, is considered by Scudder to have
several points of resemblance to Sialidae.
xt
Fia. 293. — Mormolucoides
articulatus, larva. Trias
of Connecticut. (After
Scudder.)
Fam. IX. Panorpidae—Scorpion-flies.
Head prolonged to form a deflexed beak, provided with palpi near
Fig. 294.—Panorpa communis, male,
Cambridge.
of the mouth-parts.
its apex; wings elongate and
narrow, shining and destitute of
hair, with numerous, slightly
divergent veins and moderately
numerous transverse veinlets (in
one genus the wings are absent).
Larvae provided with legs, and
usually with numerous prolegs
like the saw-flies: habits car-
NUVOTOUS.
The majority of the members
of this family are very readily
distinguished by the beak-like
front of the head, this being
chiefly due to enlargement of
parts of the head itself, and
in a less degree to prolongation
The upper (or front) face of the beak is
formed entirely by the clypeus, the labrum being scarcely
VOL. V
2G
450 . NEUROPTERA CHAP.
visible, though it may be detected at the sides of the tip of the
beak; the sutures between the various parts of the head are
nearly or quite obliterated, but it is probable that the sides of
the beak are formed by the genae and by the stipites of the .
maxillae, and its under-surface chiefly by the submentum: the
mentum itself is but small, the ligula is small, bifid.at the ex-
tremity, and each branch bears a two-jointed palpus, the
basal article being of very peculiar structure in Panorpa. The
mandibles are but small, and are placed at the apex of the beak ;
they have each the form of an oblong plate armed with two
very sharp teeth, and they cross freely. The maxillae are the
only parts of the mouth-pieces that are very elongated; each
cardo is articulated at the base of the head, and the stipes extends
all the length of the side of the beak; each maxilla bears a five-
jointed palpus and two small but very densely ciliated lobes.
The antennae are long, very slender, and flexible, and are many-
jointed; they are inserted between the eyes in large foramina ;
there are three ocelli, or none, and the compound eyes are
moderately large. The prothorax is small, its notum is quite
small or moderate in size, and the prothoracic stigma is placed
behind it; the side-pieces are small, and there is no chitinous pro-
sternum except a small longitudinal strip placed in the mem-
brane between the coxae; these latter are of only moderate size,
and are free and dependent. The meso- and meta-thorax are
large, their side-pieces are of considerable dimensions and bear
large, dependent coxae and supporting-pieces (Fig. 58); there is a —
stigma placed between the meso- and meta-thorax at the hind
margin of the upper part of the meso-trochantin ; both meso- and
meta-notum are transversely divided. The abdomen is elongate,
slender, conico-cylindrical, consisting of nine segments; the basal
segment is membranous and concealed; the terminal appendages
are of variable nature according to the species and sex. ‘The legs
are elongate and slender, the tarsi five-jointed. The internal
anatomy of Panorpa communis has been examined by Dufour?
and Loew.” They agree in describing the alimentary canal as
being of peculiar structure: there is a short, slender oesophagus
leading to an organ in which there is seated a remarkable
arrangement of elongate hairs; this structure might be looked
on as the proventriculus, but Loew considers it to be rather a
1 Mem, Ac, Sci. érang. vii. 1841, p, 582. 2 Linnaea entom, iii. 1848, p. 363.
_ 2 ee Te Eel
FA sa >
ae eee ee ee 2 ee ee
an tae ieee Le ieee =
xXx : PANORPIDAE 451
division of the true stomach. The particulars given by these
two anatomists as to some other parts of the internal anatomy
are very discrepant.
The Panorpidae form a small family of only nine or ten genera,
_ two or three of these being exotic and only imperfectly known ;
the three genera found in Europe are composed of very curious
Insects. The scorpion-flies—Panorpa proper—are very common
Insects, and have received their vernacular name from the fact
that the males have the terminal segments elongate and slender
and very mobile, and carry them curved up somewhat after the
fashion of the scorpions (Fig. 294). It is said that Aristotle was
acquainted with these Insects, and considered them to be really
winged scorpions.
A second European genus, Boreus, is still more peculiar; it is
destitute of wings, and has the appearance of a minute wingless
grasshopper; it is found
from late autumn to early
spring in moss and under
stones,andis said to be some-
times found disporting itself
on the surface of the snow :
the female of this Insect
has an exserted ovipositor.
The writer has found this
little creature in Scotland among moss in November, and under
stones early in March (Fig. 295). The third European genus,
Littacus, does not occur in our islands, but is common on many parts
of the Continent ; the perfect Insect has a great resemblance to a
Tipula, or “daddy-long-legs” fly, and attaches itself to the stems
of grasses, and preys on flies; according to Brauer it has the”
peculiar habit of using the hind pair of legs as hands (Fig. 296),
instead of the front pair, as is usual in Insects. This remark-
able genus is widely distributed, and species of it are found even
in the Antipodes. A species inhabiting caves has been mentioned
by M‘Lachlan.*
The early stages of the Panorpidae were for long unknown, but
have recently been discovered by Brauer: he obtained eggs of Panorpa
by confining a number of the perfect flies in a vessel containing
some damp earth on which was placed a piece of meat; when
1 Ent, Month. Mag. 1894, p. 39.
Fia. 295,—Boreus hiemalis, female. Dumfriesshire.
452 NEUROPTERA | CHAP,
the young larvae were hatched they buried themselves in the earth
and nourished themselves with the meat. or its juices. These
larvae (Fig. 297) bear a great resemblance to those of the Hymenop- —
terous family Tenthredinidae; they have biting mandibles and
palp-bearing maxillae, and show no approach to the peculiar
mouth structure found in the Hemerobiidae; there are three pairs
of feet placed on the three thoracic segments, and there is also a
pair of less perfect feet on each of the first eight abdominal —
segments, those behind being the larger. The upper surface of
Wallies
Fra. 296. —Bittacus tipularius holding Fra. 297.— Young larva of
a fly in its hind legs. Austria. Panorpa communis. .
(After Brauer.) (After Brauer.)
the body bears spines, which, however, disappear after the first
change of skin, with the exception of the larger processes on the
posterior segment, which persist throughout the life of the larva.
The larvae are active for about one month ; after this they become
quiescent, but do not change to the pupa state for several weeks ;
when this happens they change in form and cannot creep, although
their limbs are not enclosed in any pupa case. Brauer also dis-—
covered larvae of Panorpa communis at large in numbers in an
old tree stump that was quite covered with moss, and contained
many ants in the mouldering wood. The ants appeared to be on
friendly terms with. the Panorpa larvae. The earlier stages of
xx) PANORPIDAE—HEMEROBIIDAE 453
Boreus and LBittacus were also observed by Brauer; they are
- essentially similar to those of Panorpa, but the larva in Boreus is
not provided with abdominal prolegs. The Panorpidae have
been separated from the other Neuroptera by certain naturalists
as a distinct Order, called Panorpatae by Brauer, Mecaptera by
Packard; but in their structure as well as in their metamorphoses
they are not so distinct from the Phryganeidae and the Hemero-
biidae as to justify this step.
Fossil forms of Bittacus and of Panorpa have been fans in
amber and in the Tertiary strata, and Scudder has described some
forms from Florissant in which there are no cross-veinlets in the
wings. Some remains from the English Lias have been referred
to Panorpidae by Westwood under the name Orthophlebia, but it
is by no means certain that they really belong to the family.
(
Fam. X. Hemerobiidae—Ant-lions, Lacewing-flies, etc.
Head vertical ; macxillae free, with five-jointed palpi ;° labial palpr
three-jointed, Wings subequal in size, with much reticula-
tion, without anal area. Tarsi five-jointed. Metamorphosis
great ; the larvae with mandibles and mazxillae coadapted to
form spear-like organs that are.suctorial in function. Pupa,
similar in general form to the imago, enclosed in a cocoon.
KH -
J 7
a? :
WNL) ———
; Y i p ZED —SEA
So “ ‘Ss = are
TAY
Fic. 298.—Drepanepteryx sakes Scotland.
The Hemerobiidae are an extremely varied assemblage of
Neuroptera; the perfect Insects of the various sub-families are
very different in appearance, but the family as a whole is
naturally defined by the very peculiar structure of the mouth-
organs of the larvae. These Insects have, in fact, a suctorial
A454 NEUROPTERA CHAP.
mouth in their early life, and one of the ordinary biting type in
adult life.
This is a very unusual condition, being the reverse of what
we find in Lepidoptera and some other of the large Orders,
where the mouth is mandibulate in the young and suctorial in
the adult. The suctorial condition is in Hemerobiidae chiefly
due to modification of the mandibles; but this is never the case
in the Insects that have a suctorial mouth in the .imaginal
instar. Nearly all the Hemerobiidae are terrestrial Insects in all
their stages; a small number of them are, to a certain extent,
amphibious in the larval life, while one or two genera possess
truly aquatic larvae. The metamorphosis is, so far as the
changes of external form are concerned, quite complete. There —
are no wingless forms in the adult stage.
The classification given by Hagen’ and generally. adopted
recognises seven sub-families. These we shall mention seriatim.
Sub-Fam. 1. Myrmeleonides or Ant-lions—Antennae short,
clubbed, the apical space of the wing with regular, oblong
cellules,
paces
OS
Fic. 299.—Tomateres citrinus. 8, E. Africa. (After Hagen.)
The ant-lions in their perfect state are usually unattractive
Insects, and many are nocturnal in their habits; the species of the
genus Palpares and allies (Fig. 299) are, however, of more handsome
appearance, and attain a large expanse of wing. No member of the
sub-family is an inhabitant of Britain, though species of the typical
genus Myrmeleon are common in Central and Northern Europe. The
1 Stettin. ent. Zeit. xxvii. 1866, p. 369; this author has also sketched a classifi-
cation of the larvae in P. Boston Soc. xv. 1873, p. 248.
XX ANT-LIONS © 455
remarkable habits of their larvae attracted the attention of natur-
. alists so long ago as two hundred years. We owe to Réaumur an
accurate and interesting account of JZ formicarius, the species
found in the neighbourhood of Paris. The larvae are predaceous,
and secure their prey by means of pitfalls they excavate in the
earth, and at the bottom of which they bury themselves, leaving
only their elongate jaws projecting out of the sand at the bottom
of the pit. They move only backwards, and in forming their pit
use their broad body as a plough, and throw out the sand by
placing it on the head and then sending it to a distance with a
sudden jerk. When about to construct its trap the larva does
not commence at the centre, but makes first a circular groove of
the full circumference of the future pit. Burying its abdomen
in the surface of the earth, the Insect collects on to its head, by
means of the front leg, the sand from the side which is nearest to the
centre, and then jerks the sand to a distance. By making a second
circuit within the first one, and then another, the soil is gradu-
ally removed, and a conical pit is formed, at the bottom of which —
the ant-lion lurks, burying its body but leaving its formidable
mandibles widely extended and projecting from the sand. In this
position the young ant-lion waits patiently till some wandering
Insect trespasses on its domains. An ant or fly coming over the
edge of the pitfall finds the sand of the sloping sides yielding beneath
its body, and in its effort to secure itself probably dislodges some
more of the sand, which, descending to the bottom of the pit, brings
the lurking lion into activity. Availing himself of his power of
throwing sand with his head, the ant-lion jerks some in the
neighbourhood of the trespasser, and continues to do so until the
victim is brought to the bottom of the pit and into the very jaws
of its destroyer; then there is no further hope of escape; the
mandibles close, empale their prey, and do not relax their hold
till the body of the victim is exhausted of its juices. The position
chosen is in a place that will keep dry, as the larva cannot carry
on its operations when the sand is wet or damp, hence the soil at
the base of a high wall or a rock frequently harbours these
Insects. The parts of the mouth of the Myrmeleon are perfectly
adapted for enabling it to empty the victim without for a
‘moment relaxing its hold. There is no mouth-orifice of the
usual character, and the contents of the victim are brought
into the buccal cavity by means of a groove extending along
456 NEUROPTERA CHAP, —
the under side of each mandible; in this groove the elongate — es
and slender lobe that replaces the maxilla a
—there being no maxillary palpi—
buccal cavity at each movement a small
consisting in greater part of the two lobes
that support the labial palpi. The pharynx
is provided with a complex set of muscles,
tions as an instrument of suction. After the
, jerked away to a distance. When the —
ant-lion larva is full grown it forms a
grains of sand with fine silk from a
slender spinneret placed at the posterior
extremity of the body; in this cocoon it_
Fic. 300.—Larva of Myrme- changes to an imago of very elongate
leon pallidipennis. (After
plays backwards and forwards, probably — 2
raking or dragging backwards to the
quantity of the contents of the empaled
victim. The small lower lip is peculiar, — a
and, together with the buceal cavity, fune-
prey has been sucked dry the carcass is a
globular cocoon by fastening together a
Meinert.) form, and does not emerge until its meta-
morphosis is quite completed, the skin of the
pupa being, when the Insect emerges, left behind in the cocoon,
The names by which the European ant-lion has been known are
very numerous. It was called Formicajo.and Formicario by Vallis-
neri about two hundred years ago; Réaumur called it Formica-leo,
and this was adopted by some modern authors as a generic name
for some other of the ant-lions. The French people call these a
Insects Fourmilions, of which ant-lion is our English equivalent.
The Latinised form of the term ant-lion, Formicaleo, is not now
applied to the common ant-lion as a generic term, it having been
proposed to replace it by Myrmecoleon, Myrmeleo, or Myrmeleon ; —
this latter name at present seems likely to become generally
adopted. There are several species of the genus found in Europe,
and their trivial names have been confounded by various authors
in such a way as to make it quite uncertain, without reference to
a synonymic list, what species is intended by any particular writer.
The species found in the neighbourhood of Paris, and to which it
may be presumed Réaumur’s history refers, is now called Myrme-
XxX MYRMELEONIDES 457
leon formicarium by Hagen and others; M‘Lachlan renamed it
M. europaeus, but now considers it to be the IM. nostras of
Fourcroy. The popular name appears to be due to the fact that
~-ants—Formica in Latin, Fourmi in French—form a large part of
the victims ; while lion—the other part of the name—is doubt-
less due to its prowess as a destroyer of animal life, though, as
Réaumur long ago remarked, it is a mistake to apply the term
lion to an Insect that captures its prey by strategy and by
snares rather than by rapidity and strength. The imago of
Myrmeleon is of shy disposition, and is rarely seen even in
localities where the larva is abundant. It is of nocturnal pe
and is considered by Dufour to be carnivorous.
Considerable difference of opinion has existed as to the structure
of the mouth and of the alimentary canal in these larvae. Réaumur
was of opinion that there exists no posterior orifice to the alimentary
canal, but Dufour ridiculed this idea, and stated positively that
such an orifice undoubtedly exists. It is also usually said that
the mouth is closed by a membrane. Meinert has recently exam-
ined these points,’ and he states that the mouth is not closed by
any membrane, but is merely compressed. He finds that there is
no posterior exit from the stomach; that there is a compact mass
without any cavity between the stomach and the point where the
Malpighian tubes connect with the small intestine. The portions
of aliment that are not assimilated by the larva collect in the
stomach and are expelled as a mass, but only after the Insect has
become an imago. This peculiar excrementitious mass consists
externally of uric acid, and from its form and. appearance has been
mistaken for an egg by several naturalists. The posterior portions
of the alimentary canal are, according to Meinert, of a remark-
able nature. The small intestine is elongate, slender, and is
coiled. There are eight very long and slender Malpighian tubes ;
a pair of these have free extremities, but the other six: in the
posterior part of their course are surrounded by a common mem-
brane, and, following the course of the intestine, form ultimately
a dilated body seated on a coecum. These six Malpighian tubes
are considered to be partially, if not entirely, organs for the secre-
tion of silk for forming the cocoon, the coecum being a reservoir.
The canal terminates as a slender tube, which acts as a spinneret
and is surrounded by a sheath. A complex set of muscles com-
1 Ov. Danske Selsk. 1888, p. 43.
458 NEUROPTERA : CHAP.
pletes this remarkable spinning apparatus. The alimentary
canal of the imago has been described and
figured by Dufour’; it is very different
from that of the larva.
The ant-lion is capable of sustaining
prolonged fasts. Dufour kept specimens for
six months without any food. These In- .
sects are said to give off a peculiar ant-like
odour, due, it is thought, to their ant-
eating habits. Although no species in-
habits Great Britain, yet one is found in
Southern Sweden. Introduced specimens
get on very well in confinement in our
country,” and would probably flourish at
large for some. years if they were liber-
ated.
Although the number of known species
and genera of Myrmeleonides is consider-
able—that of the species being now
upwards of 300—the members of the
small genus Myrmeleon are the only forms
that are known to make pits of the kind
we have described. Other larvae? are
known similar in general form to the
Fic. 301.—Upper aspect of common ant-lion, but they walk forwards
prpraeay aie tf cr9 in the normal manner, and apparently
stomach ; ¢, free extremi- hunt their prey by lurking in a hidden
oa i hwo Malpis™ lace and, when a chance occurs, rush-
portion of other six tubes; ing on the victim with rapidity. Brauer
dh covcum 5 spimeret s has observed this habit in the case of
J, J, muscles for protruding vs é
its sheath ; gy, g, maxillary Dendroleon pantherinus in the Prater at
glands. (After Meinert.) Vianna:
The most remarkable forms of Myrmeleonides are contained
in the genus Palpares. We figure Tomateres citrinus (Fig. 299),
an allied genus found in Eastern Africa as far south as Natal.
These Insects have conspicuous blotches and marks on their
wings. The species of Méyrmeleon are similar in form, but are
smaller, more feeble, and less ornate in appearance.
1 Ann. Sci. étrang. vii. 1834, pl. 12. 2 M‘Lachlan, Ent. Month. Mag. ii. 1865, p. 73.
8 Redtenbacher, Denk. Ak. Wien, xlviii. 1884, p. 335.
xXx | - HEMEROBIIDAE 459
Pitfalls, formed in all probability by ant-lions, have been
noticed in the Galapagos islands and in Patagonia, though none
of the Insects forming them have been found.
Sub-Fam. 2. Ascalaphides.— Antennae elongate, with a knob at
the tip; the apical area of the wing with irregular cellules,
{/
UT
Fie. 302.—Ascalaphus coccajus. East Pyrenees.
The sub-family Ascalaphides is not represented by any species
in Britain, though ,
—
1 Rep. of the Entomologist, 1886, p. 510, Washington.
484 NEUROPTERA ' CHAP,
2
metamorphosis inasmuch as the larva, instead of lying free, con-
structs a cocoon in its case or other habitation in which to change
toa nymph. In the larvae that do not make use of a portable —
case the abdominal hooks are not essential, and are replaced —
by other organs differing much in structure, being sometimes
apparently of a sensitive nature, in other forms possibly respira-
tory. Miiller tells us of a carnivorous larva of this group in
which the anterior legs are armed with powerful forceps for pre- |
datory purposes.
The Hydroptilides comprise the most minute of the
Phryganeidae, and their species will prob-—
ably prove to be very numerous in well- —
watered tropical regions, though few have
yet been described from there. The per-
fect Insects (Fig. 320) bear an extreme
resemblance to small moths of the group
titute of respiratory filaments, and con-
struct portable cases of a variety of
forms, some resembling seeds. Miiller has
given particulars of a curious nature as
to the cases of some Brazilian Hydrop-
tilides; one species moors its dwelling
to a stone by means of a long silken
pigre 21) Pi vat, Cable, by this artifice combining safety
larva magnified; A, larva with the power of ranging over a con-
Kunin he (After siderable extent of water. In Diaulus
there is only a narrow slit at each end
of the case, but one side of it is provided with two chimneys to
permit the flow of water for respiratory purposes.
The larva of Oxyethira (Fig. 330) is a curious form, possess-
ing comparatively long legs, and a head and thorax slender in
comparison with the distended hind body. The cases are
fastened, for the purposes of pupation, to a leaf of a water-lily.
Some very curious anomalies as regards the development
of the wings exist in the Phryganeidae ; Anomalopteryx, for —
instance, has the wings quite short and useless for flight in
the male, while in the other sex they are ample; in Hnoicyla
—the curious Insect figured on p. 481, in which the larvae
are of terrestrial habits—we find the females with only rudiments
Tineidae. The larvae (Fig. 329) are des- —
ee ee ee ee ee ae ee
XXI+ - CADDIS-FLIES 485
of wings, while in Zhamastes the posterior wings are absent in
both sexes. These anomalies are |
at present quite inexplicable; and
we may here mention that we
are in complete ignorance as to
the functional importance of
many of the peculiarities of the
Phryganeidae. We do not know 4
why the mouth is reduced from
the normal state, the maxillary
palpi being, on the other hand,
extraordinarily developed ; we do
not know the importance of the
numerous spines and of the |
spurs on the legs, nor of the mye, 330.—Ozyethira costalis, A, Larva
hairs on the wings, although — * “5 vr nets Kisotick) Acat fy
these are amongst the most
characteristic of the special features of this group ‘of Insects.
Fossils—Abundant remains of Phryganeidae belonging to
the Tertiary epoch have been discovered. They are common in
amber, and it is a remarkable fact that a larval case has been
found in amber. This seems almost inexplicable, except on the
assumption that such larvae were of arboreal habits, a condition
that, at the present time, must be excessively rare, though the
terrestrial habits of Hnoicyla warrant us in believing it may
occur. -In the Tertiary Lake Basin at Colorado the remains of
Phryganeidae in the imago state are extremely abundant, so
that it is curious that but few such remains have been found in
Europe.. In Auvergne the so-called indusial limestone, which
is two or three yards thick over a wide area, is considered to be
composed chiefly of the cases of larvae of this family.
In the Mesozoic epoch some wings found in the lower Purbeck
strata are considered to be those of Phryganeidae; similar wings
have been found in the Lias, but this is the only evidence of
the existence of the family at that period except a tube, supposed
to. be a tarval case, detected in the Cretaceous of Bohemia.
Earlier than this nothing has been discovered that can be
connected with the family, so that at present the palaeontological
evidence appears unfavourable to the view held by some that
the Phryganeidae may be considered forms allied to the early
> =
™
ae
486 NEUROPTERA CHAP. XXI
conditions of the Lepidoptera. It should be noted that the
3
head in Phryganeidae is the most important part from a
systematic point of view, and that fossils have been chiefly
identified from the wings; this is a much more doubtful
character,;as the wings of the Phryganeidae have a simple
system of neuration, and in shape have nothing very charae
teristic.
Extinct Order Palaeodictyoptera.
This seems to be the fittest place to notice the existence of
some fossil remains from the Palaeozoic rocks that cannot be
fitly, or certainly, assigned to any of our existing Orders, and to
which the above name has consequently been given. These
remains consist chiefly of wings in a more or less imperfect state
of preservation, and it is therefore quite doubtful whether the
course of assigning them to a separate Order supposed to be ex-
tinct be correct. This is all the more doubtful when we recollect
that an Insect fossil, Zugereon bockingi, having the head with
mouth-parts of a Hemipterous or Dipterous nature, has been found,
the wings attached to it being such as, had they been found
separate, would have been considered to be Neuropterous, or at
any rate allied thereto. “About forty-two forms of Palaeodicty-
optera are assigned by Scudder to a section called Neuropteroidea,
and may therefore be considered to have’a special resemblance
to our Neuroptera. These Neuropteroidea comprise numerous
genera and no less than six families. Scudder’s view -is at
the best tentative, and is not very favourably received by some
entomologists. Brauer has, indeed, objected altogether to the
formation of this Order Palaeodictyoptera, and Brongniart has
published a list of the Palaeozoic Insects in which a system
of arrangement different to that of Scudder is adopted. In his
most recent work’ Brongniart assigns some of these Neur-
opteroidea to the families Platypterides and Protodonates, which
we have previously discussed. The whole subject of these
Palaeozoic Insect remains is still in its infancy, and it would
not be proper to accept any view as final that has yet been
stated, nor would it be fair to dismiss the subject as unimportant
because of the great divergence of opinion amongst the authorities
who have investigated it.
1 Insectes fossiles des temps primaires, 1893, p. 38.
ee Se ee ee
CHAPTER XXII
HYMENOPTERA——HYMENOPTERA SESSILIVENTRES—-CEPHIDAE—
ORYSSIDAE——SIRICIDAE——TENTHREDINIDAE OR SAWFLIES
Order IV. Hymenoptera.
Wings four, membranous, without scales, usually transparent, never
very large, the posterior pair smaller than the anterior ; the
cells formed by the nervures irregular in size and form, never
very numerous (less than twenty on the front, than fifteen on
the hind, wing). Mandibles conspicuous even when the other
parts of the mouth form a proboscis. The side-pieces of the
prothorax are disconnected from the pronotum and overlap
the prosternum, usually entirely concealing tt. The females
are furnished at the extremity of the body with either saw,
sting, or ovipositor ; these parts may either be withdrawn
into the body or be permanently protruded. The metamor-
phosis is great and abrupt, the chief changes being revealed
in the pupa disclosed at the last moult of the larva ; this moult
is frequently delayed till long. after growth has been com-
pleted. In the pupa the parts of the perfect Insect are seen
nearly free, each covered in a very delicate skin.
THE term Hymenoptera includes ants, bees, wasps, sawflies, and
the tribes of innumerable Ichneumon-flies. The Order is of
enormous extent, consisting even at present of tens of thousands
‘of described and named species, and yet .these are but few in
comparison with those that remain-unknown. It has good claims
to be considered the “highest” Order of Insects. Sir John
Lubbock says: “If we judge animals by their intelligence as
evinced in their actions, it is not the gorilla and the chimpanzee,
but the bee, and above all the ant, which approach nearest to
488 HYMENOPTERA CHAP.
‘
1
man.’ * The mechanical perfection of the structures of the
individuals, and the rapid and efficient manner in which their
functions are discharged, are very remarkable. In many species
of Hymenoptera the individuals have the habit of living together
in great societies, in
which the efforts of
the members are com-
bined for the support
of the whole society
and for the benefit
of a younger genera-
tion. To fit them
for this social life
the bodies of the
larger number of the
individuals are more
or less changed in
structure, so that
S Ss they become workers.
Tie he These workers are in
igo . all cases imperfect
females; besides
Fic. 331.—Bombus lucorum. A, Adult larva; B, pupa ; carrying on the
C, imago, female. Britain. ordinary work of the
society, they tend and feed the young. The duty of reproduc-
tion is restricted to a single female, called a queen, or to a small —
number of such individuals in each society. The males occupy
an unimportant position in the society, and are usually much
shorter-lived than the workers and queens. The social Hymen-
optera do not form a single zoological group, but are of three
different kinds—wasps, bees, and ants. There are numerous
non-social, or solitary, wasps and bees.
In the Order Hymenoptera—especially in the higher forms.
—the males and females are often different in appearance and
structure. In the ants, one of the social groups, the workers, or
imperfect females, are quite wingless. There are numerous other
groups in which species, not social, are found, having the females
wingless while the males have wings. In a few species there is
an apterous condition of the male, perhaps usually only as a
1 P. ent. Soc. London, 1866, p. Ixv.
XXII EXTERNAL STRUCTURE 489
dimorphic form. In the parasitic division there are species that
are apterous in both sexes. The structure of the outer skeleton,
or external part of the body, exhibits some peculiarities, the chief
of which is the detachment of the side-pieces of the prothorax and
their great development. Not less remarkable is the abstraction
of a segment from the abdomen to become, as it were, part of
the thorax; while between the first and second true segments of
the abdomen there exists a joint, or articulation, of the utmost
mechanical perfection, enabling the operations of stinging and
piercing to be executed with an accuracy that cannot be surpassed.
As a result of the detachment of the thoracic side-pieces, the
front legs and the structures connected with them are disjoined
from the notum, as shown in
Fig. 332, and act in connex-
ion with the head, while the
dorsal portion of the segment _
remains attached to the great.
thoracic mass. The head is
quite free from the thorax
and very mobile; the upper
organs of the mouth—the
labrum and the mandibles—
are not subject to modifica-
tions equal to those exhibited | :
by the maxillae and lower F!l® 332:—Tenthredo, with head fully ex-
. tended: «a, pleuron; 0%, pronotum; c¢,
lip, which parts in the bees membrane ; d, mesonotum.
are prolonged to form a suc- 7
torial apparatus that may exceed in length the whole body of
the Insect. The mandibles remain cutting or crushing imple-
ments even when the maxillae and lower lip are modified to
form a suctorial apparatus of the kind we have mentioned; so
that in the higher forms—ants, bees, and wasps—the mouth-
pieces are completely differentiated for two sets of functions, one
industrial, the other nutritive. 3
Behind the head there is a large consolidated mass represent-
ing the thorax of other Insects, but made up, as we have already
indicated, in an unusual manner, and which therefore may be
called by a special name, the alitrunk (Fig. 333). The pronotum
forms the anterior part of the alitrunk, with which it is usually
very closely connected, being indeed frequently immovably soldered
490
HYMENOPTERA
thereto.
Fic. 333.—Alitrunk of Sphex
chrysis. A, Dorsal aspect :
a, pronotum ; 0, 7, mesono-
tum; c, tegula; d, base of
anterior, ¢,of posterior, wing ;
g, division of metanotum ; A,
median (true first abdomi-
nal) segment ; 7, its spir-
acle; k, second abdominal
segment, usually called the
petiole or first abdominal
segment. B, Posterior as-
pect of the median seg-
ment: a, upper part; 34,
superior, c, inferior abdomi-
nal foramen; d, ventral
plate of median segment ;
€, COxXa.
It exhibits, however, considerable variety, and is seen
in its simplest and least soldered state in
Cephus. In the higher bees the pro-
notum takes on a form not seen in any
other Insects, being one of the most
beautiful sclerites to be found in the
class (Fig. 334, pronotum of ylocopa).
We have already remarked that in
Hymenoptera the lower portions of the
prothoracic segment are detached from
the upper, so ,
that the pro-
notum is not
supported be-
neath by a
sternum as xa
usual, In the py¢, 334.—Pronotum of a car-
bees in ques- penter bee, Xylocopa sp.
| East India.
tion the pro-
notum makes up for the removal of the
corresponding side-pieces and sternum,
by becoming itself a complete ring,
its sides being prolonged and meeting
in the middle line of the under sur-
face of the body. At the same time
a large lobe is developed laterally on
each side, overlying and protecting the
first breathing orifice. The intermediate
stages of this remarkable modification
may be observed by dissecting a small
series of genera of bees.
Although the prosternum of a Hymen-
opterous Insect is not usually visible
owing to its being overwrapped by the
CHAP.
side-pieces, it is really, as shown in Fig. 335, B, of complicated
form. In Cimbex and some other sawflies the side-pieces are not so
large as usual, but the prosternum is larger and is exposed. The
prothoracic spiracle is rarely visible externally, but its position
is remarkably constant, and is usually indicated by a peculiar
lobe or angle of the pronotum projecting backwards just below
“XXII EXTERNAL STRUCTURE 491
the insertion of the front wing. This stigmatic lobe is frequently
fringed with short hairs.
The mesothorax is the largest of the three divisions of the
thorax proper; its notum is large, and usually divided into two
parts by a transverse suture. The side-
pieces are so placed that the epimeron is
rather behind than below the episternum.
The mesosternum forms the larger part of
_the under-surface of the alitrunk. A very
large phragma projects from the meso-
thorax into the interior of the body. The
mesothoracic spiracle is usually not visible ;
its existence was unknown to the older
entomotomists, who were in consequence led
to consider the spiracle of the median seg-
ment as belonging to the thorax. The meso-
thoracic spiracle is, however, easily seen in
Cimbex, placed in the suture between the
mesothoracic epimeron and the metathoracic
Fic. 335.—Articulation of
front legs of the hornet
(Vespa crabro, 2). A:
a, side-piece of pro-
episternum, a little below the insertion of
the front wing; close to this spot the meso-
phragma, just spoken of, comes, in Cimbex,
thorax overlying the
prosternum ; 0}, coxa ;
¢, trochanter. B, pro-
sternum proper, as seen
from front when ex-
to the surface. The mesothoracic spiracle
tracted.
is generally conspicuous in the worker ant.
The parts of the metathorax are usually small, but so much variety
prevails in this respect that no general description can be given.
The structure of the posterior part of the alitrunk has given
rise to an anatomical discussion that has extended over three-
quarters of a century, with the result that it is now clear that
the posterior part of what appears to be thorax in Hymenoptera
is composed of an abdominal segment. This part has been called:
“Latreille’s segment,” the “median segment,” and the “ pro-
podeum.” The latter term was proposed by Newman, under the
form of propodeon,? and appears to be on the whole the most
1 For a history of this complex question, see Gosch, Naturhist. Tidskr. (Rk. 3)
vol. xiii. 1881 ; and also Brauer, Sitzb. Ak. Wien, \xxxv. 1882.
2 Introd. hist. Insects, 1841, p. 143. The names proposed by Newman may be
adopted when it is specially requisite to use terms that are morphologically correct.
According to his nomenclature the true whole abdomen of petiolate Hymenoptera
consists of three anatomical parts: 1, the petiole or podeon ; 2, the propodeon or
part in front of the petiole ; 3, the metapodeon or part behind the petiole.
492 HYMENOPTERA CHAP.
suitable term for this part, which is of great importance in
systematic entomology, owing to the extreme variety of characters
it affords. Although it is clear that the propodeum is, in large
part, an abdominal segment, yet its morphology is still uncertain ;
what parts are pleural, what tergal, and what may be chitinised -
spiracular area, or portions of the metathorax, being undetermined.
The ventral portion of the propodeum affords a strong contrast —
to the dorsal part, being so small that it has frequently been
described as absent; it 1s, however, not difficult to detect it in
the position shown at d, Fig. 333, B..
Although the true first segment of the abdomen is detached from
its normal position and added to the thorax, yet the term abdomen
is conventionally restricted to the part
that commences with the true second
segment, which, in counting the number
of abdominal segments, is reckoned as
being the first. There are two modes
of articulation of the Hymenopterous
abdomen with the alitrunk; in one
remains of the calibre usual in Insects,
. while in the other (Fig. 336, B) it is
Fic. 336.—Articulation of abdo-
wen, pnd altemee ee at cae greatly contracted, so that the two
hex, B, Vespa. a, Propodeum parts are connected only by a slender
or median segment; 0, dorsal : ; ;
plate Be iMae Glee se stalk, the petiole. The petiole, besides
abdominal segment or petiole; articulating in a very perfect manner
2 spate of the propodeum with the propodeum by means of cer-
, hind coxa; e, ventral plate : . : é
of first (second true) abdomi- tain prominences and notches, is also
ee connected therewith by means of a
slender ligament placed on its dorsal aspect and called the
funiculus, shown in Fig. 333, A, just at the extremity of the
pointing line % This mode of articulation gives great freedom
of motion, so that in some Petiolata (Ampules) the abdomen can
be doubled under the body and the sting brought to the head.
It is worthy of note that even in the Sessiliventres—as the sub-
Order with broad-based abdomen is called——-some amount of
movement exists at the corresponding spot; while, as shown in |
Fig. 336, A, between a and 0, there exists an exposed membrane,
the homologue of the funiculus.
The number of abdominal segments that can be seen in the
(Fig. 336, A) the base of the abdomen
XXII EXTERNAL STRUCTURE 493
perfect Insect varies greatly. In Tenthredinidae nine can be
distinguished, while in some of the Chrysididae it is difficult to
detect more than three behind the petiole. These distinctions
are, however, superficial or secondary, being due to changes in
the later life in connexion with the stings or borers; in the
larvae that have been examined thirteen segments behind the
head have usually been detected. |
Nothing is more remarkable in the Hyménoptera than the
great differences that exist in the form of the petiole. This may
be very short, as in the bees, so that the abdomen when not
deflexed does not appear to be separated from the thorax (Fig.
831, C); in this condition it is said to be sessile, a term which
it would be well to replace by that of pseudosessile. In many
of the solitary wasps the petiole is very long. In ants it is re-
placed by one or two curiously-shaped small segments called
nodes (Fig. 60, B, 2,3), and in many ants these are provided
with structures for the production of sound. The abdomen is
formed by a system of double imbrications; each dorsal plate
overlaps each ventral plate, and the hind margin of each segment
embraces the front part of the one following; thus this part of
the body has not only great mobility, but is also capable of much
distension and extension. The pleura are apparently absent, but
each one has really become a part of the dorsal plate of the seg-
ment to which it belongs. This is shown to be the case by
Cimbex, where the division between pleuron and dorsal plate
exists on each segment except the basal one. Owing to this
arrangement, the abdominal stigmata in Hymenoptera appear
to be placed on the dorsal plates.
The organs for mechanical purposes existing at the extremity -
of the body in Hymenoptera exhibit a great diversity of form ;
they are saws, borers, piercers, or stings. Notwithstanding
their great differences they are all, in their origin, essentially
similar, and consist’ of six parts developed from limb-like pro-
longations on the penultimate and antepenultimate segments of
the larva, as described by Packard and Dewitz These processes
have by some been-thought to be not essentially different from
abdominal legs, and Cholodkovsky has recently advocated this
opinion.”
1 Zeitschr. wiss. Zool. xxv. 1874, p. 184.
2 Ann. Mag. Nat. Hist. (6) x. 1892, p. 442.
494 HYMENOPTERA CHAP.
The legs of bees exhibit modifications for industrial purposes.
In the stinging Hymenoptera the trochanters are usually of a
single piece, and these Insects are called monotrochous ; but in most
of the other forms the trochanters are more or less distinctly
divided into two parts (Fig. 345, 0). The usual number of joints
in the tarsus is five, but is subject to diminution in many cases.
In the bees and ants the first joint is altered in form; in the
bees to act as an instrument for gathering or carrying pollen; in
the ants to act, as it were, as a second tibia. Between the
claws there is a very perfect pad, already described and figured
on p. 106.
The wings are remarkable for the beautiful manner in whielt
the hinder one is united to the
anterior, so that the two act in
B flight as a single organ. The
hind wing is furnished with a
series of hooks, and the hind
margin of the front wing is
curled over so that the hooks
catch on to it. In some of the
parasitic forms the wings are
almost destitute of nervures,
and have no hooks. The powers
~ of flight in these cases are prob-
ably but small, the wings merely
serving to float the Insect in
the air. In some Hymenoptera,
- especially in Pompilides and
Fic. 337.—Wings of a carpenter bee. A, Xylocopa, the wings may be
The pair of wings separated ; a, posi- deeply pigmented or even me-
tion of the hooks: B, the same wings P
when united by the hooks. C, Portions tallic; and in some forms of
of the two wings: a, the series of Tenthredinidae, Ichneumonidae,
hooks ; 6, marginal hairs ; c, portion of a ;
edge of front wing, of which the other and Braconidae the pligmenta-
part has been broken away in order to tion assumes the form of definite
show the hooks.
patterns.
The studies of the internal anatomy of Hymenoptera are at
present by no means numerous or extensive. The alimentary canal
(Fig. 69) possesses a crop, gizzard, and chylific stomach in addition
to the oesophagus and intestine. The social Hymenoptera have
the power of disgorging matter from the alimentary canal for the
wal Me a a
XXII ANATOMY 495
purpose of supplying food for their young. The crop—which
is situated in the anterior part of the abdomen—is the reser-
voir that contains this matter. The mode of disgorgement is
believed to be pressure exerted on the crop by contraction of the
abdomen. Salivary glands are present in remarkable variety.
The tracheal system possesses, in the higher winged forms, large
saccular dilatations situated at the side of the abdomen. The
nervous system is of peculiar interest on account of the high
intelligence of many of the members of this Order; and on this
point of the anatomy, Brandt’ has made rather extensive inves-
Fic. 338.— Central nervous system
. (supra-oesophageal ganglion and ven-
tral chain) of a worker ant, Cam-
ponotus ligniperdus. (After Forel.)
a, Cerebral hemisphere ; 6, primor-
dial cerebral lobe or pedunculate
body (depressed so as to show other
parts); ¢, olfactory lobe (raised
from natural position); d, nerve
to labrum; e, antennary nerve; /,
scape of antenna; g, eye; A, optic
nerve ; 7, longitudinal commissures
connecting the hidden sub-oesopha-
geal ganglion with k, the prothoracic
ganglion ; 7, mesothoracic, m, meta-
thoracic ganglion ; s, ganglion of the
petiole ; , nerve from petiole to
other part of abdomen ; 7, qg, 0, 2nd,
3rd, 4th abdominal ganglia; p, ter-
minal nerve to cloaca; ¢ bases of
legs. “
tigations, having examined it in the adult of seventy-eight
species, and in the larva of twenty-two. In the adult there are
two cephalic—the supra- and the sub-oesophageal—two or three
thoracic, and from three to seven abdominal ganglia. The bees,
wasps, and some other of the Aculeata have only two thoracic
ganglia, while some ants have three. The supra-oesophageal
ganglion is very large. The most remarkable fact revealed by
Brandt’s investigations is the. great difference that exists between
the sexes and the worker caste in the same species. The pedun-
1 OR. Ac. Paris, \xxxiii. 1876, p. 613, and Ann. Mag. Nat. Hist. (4) xviii. 1876,
p- 504; also Horae Soc. Ross. xv. 1880, pp. 20 and 31.
496 HYMENOPTERA CHAP.
culate bodies of the supra-oesophageal ganglion are considered to
be in their development correlative with that of the intelligence
or instinct. In the workers of the social Hymenoptera these
bodies are very large, while in the males and females they are
small. The workers and females of Bombus have six abdominal
ganglia, while the males have only five; and the worker of the
honey-bee has five abdominal ganglia, while the male and the
queen-bee have but four. In the leaf-cutting bee (Megachile) the
male has four abdominal ganglia and the female five, and in the
wasps the workers have five, the males and females six. The
nervous system in the larvae shows but little difference between
the ganglia, which are thirteen in number, eight being abdominal.
In the embryo of the bee Kowalewsky has observed seventeen
ganglia. The changes that take place from the embryonic
to the imago condition are therefore. dirécted to the reduction
in number of the ganglia, which is accomplished by the
fusion of some of them. In the adult Hymenopterous Insect it
would appear that the first abdominal ganglion is always joined
with the last thoracic. |
Sub-Orders.—The distinction in the form of the abdominal
articulation, previously alluded to (p. 492, Fig. 336, A, B),
divides the Hymenoptera into two great sub-Orders, the members
of which are very different in their habits and life-histories.
The Sessiliventres are plant-eaters ; their larvae (Fig. 343, A) are
provided with legs, and are able to procure their vegetable food
for themselves. The larvae of the Petiolata are maggot-like
and helpless, and are dependent for food on supplies afforded
them by their parents or companions. It is said by Dewitz that
although the larvae of the Petiolata appear to be legless, there are
thoracic legs within the body. The metamorphosis, so far as it
is known, and the early life-history of the Sessiliventres are very
similar to those of butterflies and moths, except that the pupa is
soft and has no hard external skin. A few of these plant-eating
Sessiliventres become carnivorous in the perfect state—a change
of habit that is most unusual in Insects, though the reverse
occurrence is common. The larvae of the Petiolata exhibit, in
the cases that have been examined, the peculiarity that the
alimentary canal has not any outlet posteriorly until the ter-
mination of the larval stage of existence is approaching. In
some cases there is no anal orifice; in others this orifice exists,
XXII HYMENOPTERA 497
Te
but there is no communication between the stomach and the
posterior intestine.
Packard informs us! that in Bombus the larva, after it is
full fed, passes into the pupa state (Fig. 331, A, B) by a
series of transformations accompanied by moultings of the skin.
Packard’s statements have been confirmed by others, but details
have not been fully given, so that the number of the moults,
their intervals and other particulars, are still unknown. We
have remarked that the pupal instar is very like the perfect instar,
except that it is colourless and soft, and that each of the members
is wrapped in a very delicate skin; the colour appears gradually.
This metamorphosis exhibits important differences from that of
the Lepidoptera. Packard calls the Insect, during the stages of
transformation from the full-fed larva to the pupa, the semi-pupa ;
the later stages of the pupa, when the colouring has appeared, he
terms the subimago. Altogether he considers there is a series of
at least ten moultings of the skin. His ideas were apparently
derived from examination of a series of specimens after death
rather than from observation of the development in living indi-
viduals. The parasitic forms of Hymenoptera have apparently
extraordinary metamorpheses of very varied kinds.
Parthenogenesis.—One of the most remarkable facts con-
nected with this Order is the prevalence of parthenogenesis in a
considerable number of widely separated species. In many of
these Hymenoptera it is not a mere occasional occurrence, but
plays an important part in the continuity of the species; in-
deed, it is believed that in some. members of the Order the
reproduction is entirely parthenogenetic. We shall give par-
ticulars as to some of these cases in subsequent chapters, and
will here make some remarks on the different forms of partheno-
genesis existing in the Order. The three forms of parthenogenesis
mentioned on p. 141 all occur in Hymenoptera. In the gall-
making Cynipidae parthenogenesis is frequently accompanied with
alternation of generations, a generation consisting of the two sexes
being followed by another consisting entirely of females, which
in its turn gives origin to a bisexual generation. In this case
deuterotokous parthenogenesis is established as a part of the normal
economy of the species. This same form of parthenogenesis also
occurs in other species of Cynipidae unaccompanied by alternation
1 P. Boston Soc. x. 1866, p. 279.
VOL. Vv 2K
498 HYMENOPTERA _ CHAP.
of generations. Thus in Fhodites rosae the generations resemble
one another, and the male is very rare, but is still occasionally
produced,’ and the same condition exists in other Cynipidae.
According to the observations of Adler, we may assume that the’
male, in the latter cases, is useless, the continuation of the species
being effected by virgin females although males exist. Deutero-
kous parthenogenesis also occurs.in the sawflies, but as a com-
paratively rare phenomenon.?
Thelyotokous parthenogenesis is common in sawflies, and it
also occurs in some Cynipidae. There are several species of this —
latter family in which no males have ever been found.’ The
phenomena in Lhodites rosae we have mentioned, give rise to
the idea that in that species deuterotokous parthenogenesis occurs
as an exception, the species being usually thelyotokous. Cameron, op. cit. iv. 1893, p. 9.
4 Brit. Phyt. Hym. i. p. 27. . Fletcher’s record, referred to by Cameron, men-
tions V. miliaris, but this name was probably erroneous.
a es
XXII PARTHENOGENESIS AND SEX 499
entirely of one sex, but which sex that is differs according to
other circumstances.
Production of Sex.—lIt is believed that a very peculiar form
of parthenogenesis exists in the honey-bee, and it is confidently
stated that the drones, or males, of that species are always pro-
duced from unfertilised eggs. These views are commonly called
the Dzierzon theory, and are widely accepted. They assume
that the eggs are male till fertilised, and then become female.
After the queen-bee is fertilised most of the spermatozoa soon
find their way into a small chamber, the spermatheca, near the
posterior orifice of the body; it is believed that each egg may
be fertilised as it passes the door of this chamber, and that the
eggs that produce females (i.e. workers or queens) are so ferti-
lised, but that the eggs that produce drones are not fertilised.
Hence it is supposed that the sex is determined by this act of
fertilisation, and Cheshire has described what he calls an appa-
ratus for differentiating the sexes. It is also confidently stated
that no male honey-bee ever has a father.
The facts we have stated as to the sexes resulting from
parthenogenetic reproduction in Hymenoptera generally, are
extremely opposed to the Dzierzon theory, in so far as this
relates to the production of sex. There have always been
entomologists’ who have considered this view unsatisfactory,
and the observations of several recent French naturalists* are
unfavourable to the idea that the sex of an egg is determined by
its fertilisation.
There can be no doubt that the queen honey-bee frequently
produces males parthenogenetically, and the error of the views
we are alluding to consists in taking the parthenogenesis to be
the cause of the sex of the individual. It must be recollected
that the laying of an unfertilised egg by a fertilised female may
be different physiologically from the laying of an egg by an
unfertilised female; for, though both have as result an un-
fertilised egg, it is possible that the fertilisation of the female
may initiate processes that modify the sex of the eggs produced
by the ovaries, so that though these may produce previous to
fertilisation only male eggs, yet after fertilisation they may
produce eggs of the opposite sex or of both sexes. In other
1 See Perez and Cameron, 7. Nat. Hist. Soc. Glasgow, n.s. ii.1889, p. 194.
2 Fabre, Marchal, Nicolas.
500 IIYMENOPTERA CHAP. XXII
words, the act of fertilisation may initiate a different condition
of nutrition of the ovaries, and this may determine the sex of
the eggs produced.
Polymorphism, or Castes.—The question of the causes of
the modified individuals forming the various castes of the social —
Hymenoptera has been much discussed. These individuals are
many of them very different in size and structure from-either of
their parents, and are also different in their habits and instincts.
This difficult subject is far from being completely elucidated.
In the case of the honey-bee it is well established that an egg of
the female sex can, after deposition, be made either into a queen
or a worker-bee by the mode of nutrition—using that word in
the largest sense. On the other hand, Dewitz thought that in
the case of the ant Formica rufa, the caste—whether worker or
winged female—is already determined in the Insect before leay-
ing the egg’ Weismann and others associate the caste with
‘some hypothetic rudiments they consider to exist at the pi
earliest stage of the embryonic, or oogenetic process.
Herbert Spencer says:” “Among these social Insects the sex
is determined by degree of nutrition while the egg is being
formed,” and “after an egg, predetermined as a female, has been
laid, the character of the produced Jnsect as a perfect female or
imperfect- female is determined by the nutrition of the larva.
That is, one set of differences in structure and instincts is deter-
mined by nutrition before the egg is laid, and a further set of
differences in structures and instincts 1s determined by nutrition
after the egg is laid.”
Spencer’s generalisation is not inconsistent with the facts
hitherto brought to’ light, though it is possible that the progress
of knowledge may show some variety as to the periods of the
development at which the commencements of the modifications
occur.
Fig. 339 represents the chief castes, or adult forms, existing
in a community of one of the most highly developed of the—
species of social Hymenoptera, the leaf-cutting ant, Atta cepha-
lotes. We shall, when dealing with Formicidae, enter into some
details as to these and other cases of polymorphism. Our object
0% Rajeinder to Professor Weisser; p- i. eee from Contemporary Review,
December 1893.
a ee Oe ee eS ee ee ee ee ee ee ee eee
Fig. 339.—Adult forms of Atta (Oecodoma) cephalotes, taken from a nest in Trinidad
by Mr. J. H. Hart, 25th June 1895. A, male; B, winged female; C-F, various
forms unwinged; C, so-called soldier; D, large worker; E, smaller worker; F,
smallest worker or nurse. All equally magnified (one and half times),
502 ‘ HYMENOPTERA CHAP.
at present is to bring to the eye of the reader the great diversity
of outer form that is believed, rightly or wrongly, to result from
the mode of treatment of the young. And we will also take this
opportunity of more fully illustrating the remark we made on
p. 85 as to the profound distinctions that exist between ants
and white ants, or Termites, notwithstanding the remarkable
analogies that we shall find to exist in many of their social
arrangements. | .
The analogies we allude to, coupled with the fact that there
is a certain general resemblance in outer form between the
workers of Termites and ants, and even between the extra-
ordinary castes called soldiers in the two groups, have given rise
to the idea that there is a zoological relationship between the
social forms of Neuroptera and Hymenoptera. The two are,
however, zoologically amongst the most different of Insects.
The external skeleton in Termites is remarkable for its im-
perfect development, the sclerites being small and isolated, while
the segmental differentiation of the body is low (Fig. 225, ete.),
so that there is no difficulty in counting the segments. In ants
the reverse is the case as regards both these facts, the various
segments being most unequal, so that their homologies have only
*been detected after prolonged studies, while the chitinisation and
articulation of the various parts is so complete that the ant may
be described as cased in armour, fitting together so exactly that
it is difficult anywhere to introduce the point of a needle into —
its chinks. The wings of the two kinds of Insects are also
extremely different. The differences between the modes of
growth and development of the two sets of Insects are as pro-
found as the distinctions in their anatomy. Termitidae belong
to the division of Insects in which the wings are developed
outside the body; Hymenoptera to the division’ in which they
are developed inside the body. In Termites the growth of
the individual is slow, and the final form is reached gradually.
In the ants the growth is carried on with great rapidity, and
during it the Insect is a helpless maggot absolutely dependent
on the attentions of its seniors, while the difference in form and
structure between the. ant-larva and the ant are enormous.
Both anatomy and ontogeny are profoundly different in ants
and Termites. To these distinctions must be added, as of much
importance, the fact that in Hymenoptera only the female sex
ee, Fe ee ee
XXII SUB-ORDERS *503
is modified for the division of labour, while in Termites both
sexes undergo this change. Hence it is impossible to suppose
that the remarkable analogies that exist between the societies of
ants and those of Termites are due to any common origin. It
is probably to some similar physiological susceptibilities in the
ancestors, at an extremely remote epoch, of both groups that we
must look for an explanation of the interesting resemblances in
the social lives of ants and Termites.
The Hymenoptera are no doubt one of the largest Orders of
Insécts, the species of the parasitic tribes being apparently
innumerable. No doubt 250,000 species of the Order exist; and
possibly the number may prove to be very much larger. Up to
the present time 25,000 or 30,000 have been discovered. No
remains of Insects of this Order, of older age than the Lias,
have been brought to light; it is indeed doubtful whether the
fossils considered to be Hymenopterous of the period referred
to are really such.
The Order, as already mentioned, consists of two very distinct
sub-Orders, viz. :—
1. Hymenoptera Sessiliventres.—Insects with the abdomen broad at the base,
its first segment not completely amalgamated with the thorax,
2. Hymenoptera Petioliventres or Petiolata.—The abdomen connected with
what appears to be the thorax by a slender joint, the posterior
part of the apparent thorax consisting of an abdominal segment.
Hymenoptera Sessiliventres.—This group has been variously
called Hymenoptera phytophaga, H. securifera, H. sessiliventres,
Hi. serrifera, H. symphyta. We prefer an old term, taken from
a character that enables us to recognise at a glance which group
a species belongs to. The division or sub-Order may be formally
defined as follows :— |
Abdomen nearly continuous in outline with the thorax, the two
parts having a broad connexion instead of a small highly
mobile articulation. Anal lobe of hind wings usually of
considerable size. Trochanters «ditrochous (transversely
divided into two, Fig. 345). Hatremity of body of female
furnished with saws or boring instruments, usually concealed,
in some cases visible in part. Larvae with complex mouth-
parts; three parrs of thoracic legs (imperfect in Cephidae and
504 HYMENOPTERA CHAP.
Siricidae), and frequently with numerous abdominal _ legs,
which are destitute of hooks. Food vegetable.
The Insects of this sub-Order never exhibit the highly
specialised habits and activity of the better known petiolate
Hymenoptera. Though the food in the larval stages is always
vegetable, there is considerable variety in the larvae and their
habits ; some feed in galls, some in the twigs of plants, some in
the hard wood of trees and shrubs. The majority, however, live
on the leaves of plants. Those that live in wood (Fig. 342, C)
resemble in appearance Coleopterous larvae that have similar
habits, and those that live on leaves (Fig. 343, A) resemble
Lepidopterous larvae that do likewise. There are four families
included in the sub-Order, viz. Cephidae, Oryssidae, Siricidae,
Tenthredinidae. ;
The British Sessiliventres—under the name Phytophagous
Hymenoptera—have recently been monographed by Mr. Peter
Cameron in a series of vols. published by the Ray Society.’
These contain many figures and many details relating to natural
history, in addition to the descriptions of genera and species.
Fam. I. Cephidae—Stem Sawflies.
Slender Insects, with weak integument ; free, more or less elongate
pronotum ; one spine on the front tibia. Larvae living in the
stems of plants or in the tender shoots of trees and shrubs.
The obscure little Insects composing this family have slender
antennae of peculiar form, composed of eighteen to thirty joints,
two of which are short and. stout; then come several long joints, |
with more or less power of movement, the terminal portion
consisting of an elongate club of many joints with little power
of movement. The pronotum is longer than is usual in the
Hymenoptera, and instead of being very closely connected with —
the mesonotum, it is free and mobile, although its -base over-
wraps the front of the mesonotum. The median plate (7.e. the
dorsal plate connecting the thorax and abdomen) is divided to
the base along the middle, the divisions being separated by a
membranous piece broader behind; the anal lobe of the posterior
1 Mon. Brit. Phyt. Hym. 4 vols. 1882 to 1893.
a
an§
eo eee
XXII CEPHIDAE 505
wings is small but distinct. The female bears a saw at the
extremity of the body, but it is covered by two flaps; these
form a short, terminal projection. Although too much neglected,
the Cephidae are really of great interest
as being of more imperfect or primi-
tive structure than any of the other
families of Hymenoptera. The larval
history has been traced in_ several
species. C. pygmaeus is sometimes very
injurious to corn crops on the con-
tinent of Europe, and even in our own
country its effects in this respect are
considered to be occasionally serious.
The egg is laid in the stem of the corn
plant; the larva soon hatches and eats
its way upwards in the stem. It is a soft mem a ee
grub, apparently footless, but really pos- female imago. _ Britain.
sessing six small projections in place of — After Curtis)
thoracic legs. It occupies all the summer in feeding, and when
full fed and about to prepare for its metamorphosis, it weakens
the stem by a sort of girdling process below the ear; it then
descends in the stem to near the root, where it constructs a
transparent cocoon, in which it passes the winter as a larva,
changing to a chrysalis in the month of May, and completing
its development by appearing as a perfect Insect shortly there-
after. The girdling operation is very injurious, and causes the
corn stem, when ripe or nearly so, to break in two under the
influence of a strong wind, so that the ears fall to the ground.
The history of C. integer has been given by Riley. This
Insect attacks the young shoots of willows in North America.
Riley states’ that by a wonderful instinct the female, after she
has consigned her ege to the twig, girdles the latter, preventing
it from growing any further, and from crushing the egg. by so
doing. The larva after hatching eats downwards, sometimes
destroying-a length of two feet of the twig; when full grown it
fills the bottom of the burrow with frass, and then previous to
making its cocoon eats a passage through the side of the shoot
about a quarter of an inch above the spot where the cocoon will
be placed, thus making it easy for the perfect Insect to effect its
1 Insect Life, i. 1888, p. 8.
506 IIYMENOPTERA | CHAP.
escape ; it leaves the bark, however, untouched, and is thus pro-
tected in its retreat. A delicate transparent cocoon is then spun
in which the larva passes the winter, changing to a pupa in the
following March, and emerging as a perfect Insect about six
weeks thereafter.
Somewhat less than 100 species of this family are at present
known; the great majority are found in the Mediterranean —
region, but there are several in North America. As a single species”
is known from Mexico and another from Japan, it is probable
that the family may prove to have a wider geographical exten-
sion than at present appears to be the case.
Fam. II. Oryssidae.
The median plate behind the metanotum entire, not divided in the —
middle; antennae inserted below the eyes immediately above
the mandibles, under a sharp edge.
This family consists of the genus Oryssus, and includes only
about twenty species, but is nevertheless very widely distributed
| over the world. They are very
rare Insects, and little is known
as to their habits; one species,
O. abietinus, was formerly found
in England. Should any one
be so fortunate as to meet with
it, he can scarcely fail to re-
cognise it on noticing the
peculiar situation of the base
of the antennae. In this re-
spect the Chrysididae somewhat
resemble Oryssus, but in that
group of Hymenoptera the hind
body or abdomen is remarkably
mobile, so that the Insects can
coil themselves up by bending
Fic, 341.—Oryssus sayi. North America. at this joint ; whereas in Oryssus
A, The female Insect; B, head seen the hind body is very closely
Reece amalgamated with the thorax—
more so, in fact, than in any other Hymenopterous Insect—and
has no power of independent movement.
XXII : HYMENOPTERA 507
Oryssus abietinus very closely resembles C. sayi (Fig. 341);
it has indeed been recently suggested by Mr. Harrington that
the two supposed species may really be identical.
Fam. III. Siricidae or Uroceridae.
Pronotum closely connected with the mesonotum, perpendicular in.
JSront ; the anterior lobe of the latter not separated by the
lateral lobes from the posterior lobe: the median plate (behind
the metathorax) is divided longitudinally along the middle.
The female is provided at the extremity of the body with an
elongate, cylindrical boring instrument. The larvae live in
the wood of trees.
Fic. 342.— Zremex
columba. North
Americu A,
Imago, female:
B, pupa, female,
ventral aspect :
C, larva ; a, im-
perfect legs: D,
parasitic larva of
Thalessa. (B
and D_ after
Riley.)
The Insects of this family are usually of large size and of
bright conspicuous colours; these, however, frequently differ
greatly in the sexes of the same species, and may be very vari-
able even in one sex. The antennae are filiform and usually
elongate ; the head is usually contiguous with the thorax, but in
one division, Xyphidriides, it is exserted and separated from the
thorax by a well-marked neck. The pronotum is attached to
the mesonotum, and possesses very little, if any, freedom of
movement; it varies. in its size, being sometimes conspicuous
508 HYMENOPTERA CHAP. ~
from above; in the Xyphidriides it is smaller, and in the middle —
is entirely vertical in its direction. The mesonotum is moderate
in size, and its divisions are delimited by broad vague depressions.
The prosternum appears to be entirely membranous, but the
prosternal plates (pleura) are large, and meet together accurately
in the middle, so as to protect the greater part of the under-
surface of the neck. The abdomen is cylindrical or somewhat.
flattened above; it has seven dorsal plates in addition to the
spine-bearing terminal segment. The trochanters are double,
the outer division being, however, short; the anterior tibia has
only one spur; the anal lobe of the posterior wings is large.
The “ borer” or ovipositor of the female is a remarkable organ ;
it is held projecting directly backwards from the extremity of
the body, and has the appearance of being a powerful sting. The
apparatus is much longer than it appears, for it proceeds not
from the apex of the body, but from.the under-surface far for-
wards, so that the part exposed is only about one-half of the total
length ; it consists of a pair of elongate sheaths, which are easily
separable though they wrap together, and enclose a slender tube.
This tube is rigid and quite straight; though appearing solid, it
is really composed of two very perfectly adjusted laminae and a
third arched piece or roof. The two lower laminae are called
the spiculae; they aré serrated or grooved in a peculiar manner
near the tip, and although ‘so closely adjusted to the borer or
upper piece of the tube as to appear to form one solid whole
with it, they are said to be capable of separate motion. In
addition to these parts, the termination of the abdomen bears
above a shorter piece that projects in a parallel plane, and forms
a sort of thick spine above the ventral pieces we have described ;
this process is very strong, and has in the middle of its under-
face in Sirex gigas a membranous cavity, replaced in S. juvencus,
according to Westwood, by a pair of minute pilose styles. The
Insect, by means of this powerful apparatus, is enabled to deposit
her eggs in the solid wood of trees, in which the larva sometimes
penetrates to the depth of eight inches.
Sirex gigas is one of the most remarkable of our British
Insects, but is little known except to entomologists, being usually
rare. On the continent of Europe it is, however, an abundant
Insect, especially in the neighbourhood of forests of fir-trees,
and is a cause of considerable terror. As the Insect is not
ee ee eee te Te ey
a
»
a
=k
ee ey ae
ee ee ee ee ee
_ XXII SIRICIDAE 509
capable of inflicting much injury to the person, it is probable that
the peculiar ovipositor is believed to be a sting. The eggs are
laid—it is said to the number of 100—in the solid wood of fir-
trees, but not in perfectly healthy wood; the reason for this, it
is thought, being that in a healthy tree the great affluence of sap
caused by the burrows and presence of the Insect would be in-
jurious to the latter. The Strea will, however, attack a perfectly
healthy tree immediately after it has been felled. The larva, small
at first, enlarges its burrows as itself grows larger, and thus the
wood of a tree may be rendered completely useless for trade pur-
poses, although there may be’ very little outward indication of
unsoundness. The larva (Fig. 342, C, larva of Zremesz) is a pallid,
maggot-like creature, with six projections representing thoracic
legs; there are no other legs behind these, but some slight pro-
tuberances take their place; the terminal segment is enlarged, and
bears a hard spine. There is a difference of opinion as to the
duration of the life of the larva, Kollar saying that in seven
weeks after the deposition of the egg the maggot is full fed,.
while others consider that it takes two years to attain this con-
dition; the latter statement is more probably correct, it being
the rule that the life of wood-feeding larvae is more than usually
prolonged. After becoming full fed, the Insect may still pass a
prolonged period in the wood before emerging as a perfect Insect.
As a result of this it not infrequently happens that the Insect
emerges from wood that has been carried to a distance, and used
for buildings or for furniture. A case is recorded in which large
numbers of a species of Sivex emerged in a house in this country
some years after it was built, to the great terror of the inhabi-
tants. The wood in this case was supposed te have been brought
from Canada.
Fabre has studied! the habits of the larva of Sirex augur,
and finds that it forms tortuous galleries in the direction of the
longitudinal axis of the tree or limb, and undergoes its meta-
morphosis in the interior, leaving to the perfect Insect. the task
of finding its way out; this the creature does, not by retracing
its path along the gallery formed by the larva, but by driving a
fresh one at right angles to the previous course, thus selecting
the shortest way to freedom. By what perception or sense it
selects the road to the exterior is quite unknown. Fabre is not
1 Souvenirs entomologiques: quatriéme série, 1891, p. 308.
510 HYMENOPTERA CHAP,
able to suggest any sort of perception that might enable the larva
to pursue the right course, and considers it must be accomplished
by means of some sensibility we do not possess. Fabre’s observa-
tion is the opposite of what has been recorded in the case of 8.
gigas, where the larva is said to prepare the way for the exit of
the perfect Insect. |
Individuals of Sivex are often found in dried and solid wood,
encased by metal. When the Insect finds itself so confined; it
gnaws its way through the metal, if this be lead, and escapes.
The perseverance displayed by the Insect in these circumstances
seems to indicate a knowledge of the direction in which liberty
is to be found.
About 100 species of Siricidae are known. They form two
sub-families :—
1. Stricides: back of head nearly or quite contiguous with the pronotum.
2. Xyphidrvides: back of head separated from the pronotum by an elonga
neck. $
We are reputed to possess in Britain two species of each of
these sub-families, but it is doubtful whether more than one
Siricid is truly native. Sirex gigas is frequently brought over
in timber, and certainly breeds at times freely in Britain. Mr.
Leech has recorded the occurrence of the larvae in abundance
in fir-trees in the neighbourhood of Dublin. Sirex juvencus
is more rarely met with. Xyphidria camelus is doubtless a
native, though now apparently rare. It used to occur about eld
willows, near London, in the New Forest, and, I believe, also
in the neighbourhood of Cambridge.
Fam. IV. Tenthredinidae—Sawflies.
Hymenoptera Sessiliventres, having the pronotum small, accurately
adapted to the mesonotum ; the anterior lobe of the latter is
widely separated from the posterior; there are two spurs on
the anterior tibiae. The larvae usually live on leaves after
the manner of caterpillars, but a few inhabit galls.
The sawflies are an important family of Insects, their species
being numerous, while some of them are, in the larval state, very
destructive to vegetables and fruit. Being quiet creatures, rarely
seen on the wing, they are, though common Insects in this
ra
a fk ee ee ee
mite | Lat
XXII SAWFLIES 511
country, but little known, and few persons recognise a sawfly as
such. They are usually of small or moderate size, and the
numerous species have a great family resemblance. This remark
requires some qualification in the case of the Cimbicides, they
being Insects of larger size—usually surpassing the honey-bee—
of more robust structure,
and with greater powers of
flight.
The antennae are re-
markably variable in form
and structure. Cameron
considers that nine should
be taken as the normal
number of their joints;
but there are only three
in Hylotoma, while in Lyda
there may be forty or more.
The head is usually held
closely applied to the
thorax, but is really borne
Fic. 343.—Lophyrus pini. Britain: A, Larva; 00 @ neck capable of much
B, ventral aspect of pupa; C, imago, male- elongation (Fig. 332).
(After Vollenhoven.)
The pronotum forms .a
part of the alitrunk, but is not soldered thereto. Usually the
prosternun is more or less completely concealed by the side-
pieces, but in Cimbicides it is larger and conspicuous, the side-
pieces being in this group smaller than -usual. The dorsal
pieces of the mesothorax have their relative proportions different
to what we find them in the other families of Sessiliventres, and
even in most of the other Hymenoptera. There is first an
antero-median lobe of triangular shape projecting, like a wedge,
far backwards, into the great lateral lobes. ‘These latter form
the larger part of the area of the mesonotum ; they meet together
in the middle line, and behind are separated by a deep depression
from the posterior lobe, or scutellum of the mesothorax, which
is frequently divided into two parts, the anterior being the so-
called scutum. The pieces of the metanotum are short and
obscure, owing to the great unevenness of their parts; on each
side of the middle there is a small membranous space of pallid
colour. The cenchri, as these spaces are called, are, in Lyda,
512 HYMENOPTERA CHAP. .
delicate, membranous, depressed spaces, in front of each of which .
there stands up a flap of membrane. The function of the cenchri
is quite unknown. The median plate is fastened to the hind -
margin of the metanotum, and looks quite like one of the dorsal
plates of the following abdominal segments, from which, however,
it is separated by a more or Jess conspicuous membrane. In
the majority of the Tenthredinidae the median plate is divided
along the middle, but in the Cimbicides this is not the case.
The mesosternum is very large, and the metasternum small,
so that the middle‘and hinder pairs of coxae are placed close
together. The abdomen consists of nine segments, there being
eight dorsal plates in addition to the median plate, and seven
ventral plates besides the terminal armature. There is a pair of
short cerci, each of a single segment. The trochanters are
divided; each tibia bears two spurs at the extremity, and the
tarsi are 5-jointed.
The most characteristic and interesting of the structures with
which the Insects of this family are provided is the apparatus
from which the name of sawfly is derived. As long as two
centuries ago these instru-
ments excited the admira-
tion of Vallisnieri and of
Réaumur, who deseribed
them at length; and it is
truly astonishing that any
part of a living being
should be changed into
tools so mechanically per-
fect as these saws are
(Fig. 344). They serve
the purpose of assisting
the. female in depositing
the eggs in a suitable
situation, the place selected
being frequently the ten-
der stems of shrubs or
pair spread out and placed in a horizontal posi- ;
tion ; a, the lower margin of the saw proper; other plants, or the in-
b, the upper margin of the support: B, two
teeth of the saw more highly magnified. terior of leaves. These
organs are therefore of
course possessed only by the female. They are placed on the
XXII : SAWFLIES 513
lower aspect of the hinder extremity of the body, where they
are enclosed and protected by a pair. of sheaths, from which
they can be made to protrude by a little pressure exercised on
the parts immediately in front of them. Each female possesses
a pair of these saws; they consist of thin laminae of very
hard consistence, and are not only toothed at their edge, but in
many cases each tooth is itself serrate; at the same time the
outer face of the saw is sculptured or plicate in a remarkable
inanner, so that the saw in this way acts as a file or rasp. The
Insect having selected a suitable place, uses the saws by placing
the extremity of the abdomen against a twig or leaf, protruding
the blades, which, moving with an alternate motion, one being
thrust forward while the other is retracted, act on the plant so
as to make an incision. Each saw is directed in its movement by
the support, the pair of supports being united at the base by
- membrane as shown in Fig. 344. In the case of some species,
—Hylotoma rosae, the common sawfly of our rose-bushes, for
instance—there is no difficulty in observing the operation ; in-
deed old Réaumur, when speaking of the placid disposition of the
sawflies, suggests that it was given them so that we may easily
observe their charming operations. We cannot but regret that
in these days we are unable to take so complacent a view of the
arrangements of. nature. There is much variety in the details
of the structure of these saws; so much indeed that it is possible —
to identify most of the species by means of the saw alone.
According to certain observers, the eggs are laid by some kinds
on, not in, the leaves, so that we may conclude that in, these
cases the saws are not used by their possessors. An_ incision
having been made, an egg is placed in it, and also a drop of some
liquid matter. The egg is at first small, but soon increases till
it becomes twice or three times its former size, and the develop-
ment of the embryo commences.
The larvae of the Tenthredinidae exhibit great variety, and
are indeed in this respect more interesting than the perfect
Insects. The usual rule is that the larvae much resembles
those of Lepidopterous Insects, and feed exposed on plants in
the same way as Lepidopterous larvae do. But the exceptions
are numerous; sometimes the larva is covered with slime, and’
thus protected from various enemies. In other cases it is very
depressed, a broad creature, of irregular outline, living closely
VOL. V 21
514 , HYMENOPTERA CHAP.
attached to the leaf, somewhat after the fashion of a huge scale-
Insect. Some larvae mine between the layers of a leaf, others roll
up leaves; a few live in the stems of plants, and one or two inside
fruits. Even this does not complete the list of their habits, for
a few species of Vematus live in galls caused by the deposition of
the egg. ™:)
and are placed in a triangular position on the vertex. The pro-
notum is small in front; and extends backwards at the sides to
the points of insertion of the front wings; it is fixed to the
mesonotum. The wings (Fig. 367, A) have a more complex
neuration than those of most of the other parasitic Hymenoptera,
but are occasionally absent in one or both sexes of a species. The
metathorax is very small, and the middle and hind legs are
placed close together. The propodeum is very large, and is
552 HYMENOPTERA CHAP. |
frequently covered with a highly-developed sculpture. The hind
body springs from the lower part of the propodeum; it is usually
of slender form, and its segmentation is very conspicuous. The
females bear an ovipositor, which differs greatly in length accord-
ing to the species, and is known in the case of one species. to
attain a length six times that of the whole of the rest of the
body The egg is deposited by some species on the skin, by
others within the body of the victim ; it varies much in form and
colour, some eggs being stalked and of peculiar shape. The
larvae issuing from the eggs are legless maggots with a delicate
integument of pallid white or creamy colour. If the eggs are laid
on the surface of the body, the result-
ing larvae (except in the cases of the
interior of their victim, and disappear
therein. The changes that take place
) in the lifetime of the larvae have
been studied in only a few cases; but
if we can judge from Ratzeburg’s
history? of the changes that take
place in Anomalon, they are of great
interest. From observation of the
number of larvae of A. cirewmflecum
he distinguished four stages. It is of
course impossible to follow directly
the growth of one individual, be-
cause it is concealed in the interior
and to open this involves the death
, ; of both caterpillar and Ichneumon-
Fra. 361.— Anomalon circum- larva, The life history must therefore
Jlecum, larval development.
(After Ratzeburg.) A, First be constructed from a great number of
instar ; B, second instar ; ©, the separate observations; and it is not —
larva in the third or encysted
stage extracted from its cyst; ascertained that the four instars de- —
scribed by Ratzeburg represent the —
D, the mature larva ; E, pupa.
number of moults of the larva that actually take place. He,
however, entertained no doubt that all the forms he observed
1 Tosquinet, Ann. Soc. ent. Belgique, xxxviii. 1894, p. 694. -
2 Ichnewm. Forst. Ins. 1844, p. 81.
external parasites) soon bore into the ©
differences existing amongst a great
of the caterpillar in which it lives,
— ts? Pun
~
XXIII ICHNEUMON-FLIES 3 553
were stages in the development of one species. In the earliest
stage, when only one millimetre in length and about as thick
as a horse-hair, the larva is free in the interior of the cater-
pillar’s body, and has a small head armed only with a pair of
mandibles. There are, in addition to the head, thirteen segments,
and the last of these is an elongate tail forming nearly one-
half the length of the creature. No trace of tracheae can be
discovered. In the second stage the larva is still free, an elon-
gate tracheal tube exists, the tail has diminished to half the
length, the head has become much larger, and rudimentary
antennae of one joint are visible; possibly stigmata are present
at this stage, though they cannot afterwards be detected. In
the third stage (Fig. 361, C) the larva is encysted, the head is
large, the parts of the mouth are all developed, the tracheal
system is extensive, and the caudal termination of the body is quite
short ; notwithstanding the extensive development of the tracheal
system, no stigmata can be found. In the fourth stage the
larva is still encysted, the tail has disappeared, the head and
mouth parts are reduced in size and development, and the creature
has now the appearance of a normal larva. The changes to pupa
and perfect Insect take place within the body of the victim, in
some cases, if not usually; after it has undergone its metamor-
phosis into a chrysalis. Very little information is extant as to
the duration of the various stages, but it appears to be the rule
that only one generation appears annually, though in some cases
there are pretty certainly two.
It is very difficult to observe the. act of oviposition; the
Ichneumon-flies usually decline to notice caterpillars with which
they are placed in confinement. Ratzeburg thinks they will only
attack caterpillars that are in a deficient state of health or vitality.
Occasionally we may by a happy chance observe the act in
Insects at large, and from the records of observers it may be
deduced with tolerable certainty that the sense of sight takes no
part in the operation. Ratzeburg relates that he saw a Pimpla
alight on a leaf of Rhus and thrust its ovipositor through the
leaf. On looking to the under-side of the leaf he found that a
cocoon of Bombyx neustria was concealed there in such a position
that it could not have been seen by the Ichneumon.
Among the most remarkable of the Ichneumon-flies are the
Insects of the genera Rhyssa and Thalessa. These fine Insects
554 HYMENOPTERA CHAP.
have an ovipositor three or four inches in length, and are parasitic
on species of the family Siricidae, which, as we have previously
described, live in solid wood. In order therefore to deposit
the egg in a suitable place, the wood must be pierced by the
Ichneumon. The ovipositor is not only of extreme length, but is
also furnished with serrations on its apical part, so that it forms a —
very effective boring apparatus. It is brought into use by being
bent on itself over the back of the Insect (Fig. 362), so as to bring
the tip vertically down on to the wood, through which it is then
forced by a series of efforts; the sheaths do not enter the wood.
The egg is laid anywhere in the burrow of the Sirex ; the young
larva seeks its prey, and lives on it as an external parasite (Fig.
342,D). Erne, however, states’ that the young larva of Rhyssa
persuasoria enters its victim, and remains within the latter till its
death occurs. This happens
when the young Rhyssa is —
two or three lines in length,
and it then makes its exit from
the interior of the body and
gradually eats it up. Should
the larva it has attacked be of
large size, it of itself affords
sufficient food for the comple-
tion of the growth of the
Rhyssa. Should the Rhyssa,
however, have attacked a small
larva, this does not furnish it
« With sufficient food, and it con-
sequently dies without seeking
; Pi another larva. Erne says,
Fra. See aie Re Oviposition. Sndeod,| that at will Gees ie
another if offered to it, so that
in order to rear the Rhyssa in captivity, the victim it has first
attacked must always be given to it. The same observer states
that the Rhyssa larva is sometimes transported by the Sirex
deep into the wood, so that when it has completed its metamor-
phoses the Ichneumon-fly may find itself buried in solid wood to
a depth of about two inches.- In that case it excavates the wood
with its mandibles, and should it fail to gain the exterior after
1 Mitt. schweizer. ent. Ges. iv. 1876, p. 518,
XXIII ICHNEUMON-FLIES 555
three days of work, it dies. In the case of Zhalessa it is stated
that it sometimes bores into wood where there are no larvae, but
Riley thinks this erroneous; it is, on the other hand, certain
that the Insect after penetrating the wood is frequently unable
to withdraw the ovipositor, and consequently dies.
Packard has recorded, without mentioning the species, the
oviposition of an Ichneumon of which the egg is deposited
externally. It was placed on the head of the caterpillar, and
speedily hatched; the young larva at once bored through the
prothoracic slack of the victim, the head of the latter then
became swollen, and covered the opening into the prothorax, made
by the parasite.
The history of an Ichneumon larva that feeds as an external
parasite has been sketched by De Geer and Newport. The
observations of the latter? refer to Paniscus virgatus; he
found small, shining, black bodies attached to the skin of the
larva of a moth, Mamestra pisi; these were the eggs of the
Ichneumon. They are furnished with a short peduncle, which
is implanted in the skin of the victim; the egg, according to De
Geer, being retained more firmly by the peduncle subsequently
swelling, so as to form two knobs. The hatching takes
place by the egg-shell splitting longitudinally, while from the
split protrudes: the little head of the destroying larva. This
becomes fixed to the caterpillar, from which the nutriment is
to be drawn; the Paniscus larva does not, however, leave the
egg-shell, but, on the contrary,
becomes adherent to it, so that
the parasite is in this manner
fixed by the two ends to its
victim. In fifteen days the
parasite jessy full-grown, and had Fie. 363.—Young larva of Paniscus in
become half an inch in length. position of feeding on the skin of Mam-
At first no tracheae were ea (After Newport.) a, The egg-
to be seen, but these were
-- detected after the second day. Moulting took place three
times, and in a peculiar manner, very different from that.
described by Ratzeburg as occurring in the internal parasites
(which, he states, change their very delicate skin by detaching it
in almost imperceptible fragments). In the external parasite the
1 Fifth Rep. U.S. Ent. Comm. 1890, p. 15. 2 Tr. Linn. Soc. xxi. 1852, p. 71.
556 HYMENOPTERA CHAP.
skin remains entire, and is shuffled down to the extremity of the
body, but cannot be completely detached owing to the anchoring
of the posterior part of the body to the caterpillar; the cast
skins thus remain as envelopes to the posterior part of the body.
Newport states that if the mouth of the parasite be detached, it
usually cannot again seize hold of the victim, and consequently
perishes. It is a curious fact that more eggs than one caterpillar
can support are habitually placed on it, and some of the resulting
larvae of necessity perish during the period of growth. Poulton,
who has recently made some additional observations on the
development of Paniscus} says that if three larvae are close
together, it is the middle one that perishes, and suggests
that this is due to some simple physical condition. From
Newport’s account it may be gathered that the Mamestra
retains sufficient vitality to form its cocoon, and that the —
Paniscus larvae likewise construct their own cocoons within
that of the Mamestra. In the case of Paniscus cephalotes
feeding on Dicranura vinula, Poulton relates that the latter died
after the twelfth day of attack. The parasites, having relaxed
their hold on the victim just previous to this event, then thrust
their heads into the dead body, and devoured the larva, leaving
only a dried and empty integument. These larvae span a loose
sort of web in which to undergo their metamorphosis. In a
natural state, however, they form cocoons inside the cocoon of
the Dicranura. The period passed in the pupal condition was
about four weeks. This parasite only attacks the Lepidopterous
larva during the last stage of its existence as a larva, but the
eggs may be laid on the victim in an earlier stage; and in such
ease De Geer has stated, and Poulton has confirmed the observa-
tion, that though the larva sheds its skin it does not get rid of
the eggs.
The little Ichneumons of the genus Pezomachus are quite
destitute of wings and somewhat resemble ants; they are
common Insects in Britain. Only the female sex is known, and
it is believed that the winged Ichneumons assigned to the genus
Hemiteles—of which no females are known—are the males of
Pezomachus. Repeated efforts have been made to place this
beyond doubt, but they have usually failed, for when a brood of
these parasites is reared, the individuals generally prove to be
' Tr. ent. Soc. London, 1886, p. 162, and 1887, p. 303.
.’ ‘
scares SC ae
a Te vaee a) ee eee ee ee ee eee ee
~
XXIII ICHNEUMON-FLIES | 557
either all Hemiteles or all Pezomachus. It is to be hoped that
this interesting case will be fully elucidated.
Although the Ichneumonidae are perhaps the most purely
carnivorous of all the great families of Hymenoptera, there is
nevertheless reason for supposing that some of them can be
nourished with vegetable substances during a part at any rate of
_ the larval existence, Giraud and Cameron * having recorded observa-
tions that lead to the conclusion that some species of the genus
Pimpla may inhabit galls and live on the substance, or juices
thereof.
Over 1200 species of Ichneumonidae are known to inhabit
Britain, and there can be no doubt that this number will be
increased as a result of further observation. Unfortunately no
general work has yet been published on this department of our
fauna, and the literature is very scattered.2 The species of
North America have not received so much: investigation as those
of Europe, and the Ichneumon fauna of the tropics remains almost
uninvestigated. Six sub-families are recognised: Agriotypides,
Ichneumonides, Cryptides, Tryphonides, Pimplides, Ophionides.
Of these the first is the most remarkable, as it consists of an
Insect having aquatic habits. It
has for long been known that the
unique species Agriotypus armatus, a
rare Insect in our islands, is in the
habit of going under water and re-
maining there for a _ considerable
period, and it has now been satis-
factorily ascertained that it does this
for the purpose of laying its eggs in
the larvae of Trichoptera.® The re-
sultant larva lives inside the cases
of species of Stilo, Goéra, etc. It eet htt 5 dager Curtis)
undergoes a sort of hypermetamor-
phosis, as its shape before assuming the pupal condition
1 Ent. Month. Mag. xiii. 1877, p. 200.
2 A catalogue, with references, of the British Ichneumonidae was published by
the Entomological Society of London in 1872. Since then many additional species
have been detected and recorded, by Mr. Bridgman and others, in the Z’7ransactions
of the same Society.
8 Klapalek, Ent. Month. Mag. xxv. 1889, p, 339, and Arch. Landesdurchforschung
Bohmen, viii. No. 6, 1893, p. 53.
558 HYMENOPTERA CHAP.
is very different to what it was previously. It changes to
a pupa inside the case of the Trichopteron in a cocoon attached
to the walls. of the case. Previous to making this, however, the
Agriotypus forms a curious, elongate, string-like process attached
to the anterior extremity of its cocoon. The use of this is
unknown. Full information as to the life-history of this aquatic
Hymenopterous larva, especially as to its respiratory functions, 7
Fic. 365.—Metamorphosis of Agriotypus. (After Klapalek.) A, Larva ; B, sub-nymph ;
C, case of the Silo with the string of attachment formed by Agriotypus ; D, section
of the case: v', operculum of case ; v’, cocoon ; ag, pupa of Agriotypus ; e, exuvia
of Agriotypus ; w?, wall of cocoon ; s, remains of Silo; w', closure of case.
would be of great interest. The affinities of this remarkable
Insect are still doubtful. It may probably prove to be between
Proctotrypidae and Ichneumonidae.
Remains of Insects that may be referred with more or less
certainty to Ichneumonidae have been found in some abundance
in various tertiary strata both in Europe and North America, but
nothing indicative of the existence of the family has yet been
found in the older rocks.
Fam. V. Braconidae—Supplementary Ichneumon-flies.
Antennae with many (nearly always more than fifteen) joints, not
geniculate. Wings with a moderate number of cells, which
on the anal part of the front wing are more or less imper-
fect, the anal (i.e. the second posterior) cell being separated
from the cubital cells by a large space in which there is no
cross-nervure. Abdomen with but little mobility between the
segments ; the suture between the second and third usually
i
t ice eis a ees
r
—, eee ee ee
eg ee ao
|
:
}
}
XXIII BRACONDIDAE 559
absent, or obsolete. Larvae living parasitically in—possibly
exceptionally outside—the bodies of larvae or pupae of Insects.
Y i}
—< i <
4 (= “
——
wi
\/
Fic. 366.— Bracon palpebrator, Fic. 367.—Diagram of wing of Ichneu-
female. Europe. (After Ratze- - monid (A) and of Braconid (B). 1, 2,
burg. ) 3, 4, series of cells extending across
the wing; a, 0, divided cell of the
Ichneumonid wing, corresponding with
a, the undivided cell of the Braconid
wing.
The Braconidae are the Ichneumones, or Ichneumonides,
adsciti of the older Hymenopterists. They are extremely similar
to the Ichneumonidae, but the hind body has a much less degree
of mobility of its segments, and there are some constant distinc-
tions in the wings. Although there is a great deal of difference
in the various forms of each of the two families, yet there are
two points of distinction easily appreciated; the series of cells
running across the wing (Fig. 367) being only three in the
Ichneumonides (Fig. 367, A), but four in the Braconids (Fig. 367,
B); besides this the space a of the Braconid wing is divided
into two (a, b) in the Ichneumonid wing. A glance at these
characters enables us at once to separate correctly the thousands
of species of the two families.
The habits of the Braconidae are similar to those of Ichneu-
monidae, it being believed that all are parasites. Usually they
attack larvae, but they are bred in great numbers from pupae,
and even from imagos of other Insects. Hlasmosoma is one
of the few parasites. known to attack ants. As many as 1200
specimens of Microgaster have been reared from a single Lepi-
dopterous larva. Although such parasitism raises a feeling of
repulsion, yet there is reason for supposing that there may be
little or no cruelty or acute suffering connected with this mode of
life. The victim attacked is not eaten, the parasites in the in-
terior taking in the lymph of the caterpillar either by the mouth
or by endosmosis, but not biting their host. The latter displays
no sign of sickness, but eats voraciously, so that it serves merely
as a sort of intermediary between the juices of the plant and the
larvae inside itself. It is only when the metamorphosis is at
hand that the host sickens, but this does not always happen:
parasitised larvae frequently change to pupae, and they may
occasionally even become perfect Insects. Cases are known in
which imagos have appeared with some of the small parasites
embedded in some of the outer parts of their bodies. These
cases are, however, very rare; in the enormous majority of
instances the host is destroyed either when it is in the larval
stage or before the pupa has. advanced to any great extent on
its metamorphosis to an imago. Particulars as to various species
will be found in the valuable work of Ratzeburg we have already
referred to. Reference may also be made to Goureau’s account
of Microgaster globatus, this latter including some suggestions
by Dr. Boisduval on some of the difficult physiological questions
involved in the lives of these parasites.
The metamorphosis of Microgaster fulvipes has been studied
by Ratzeburg, and an epitome of his observations is given by
Marshall.? The larva goes through a series
of changes somewhat similar to those we
have already sketched in Anomalon cir-
cumflecum. Usually these Insects after
emerging from the body of their host spin
connected together. A most curious case
- has, however, been recorded by Marshall ®
Fic. 868.—Stalked cocoon formed as an exceptional act by Apanteles
of Apanteles formosus. a
(After Marshall.) Sormosus. Mr. Marshall has recently re
ceived other specimens of this cocoon as
well as the Insects reared therefrom in France, and inclines to
the opinion that the stalked cocoon may be the usual form, and
is sometimes departed from by the Insect for unknown reasons.
1 Ichnewm. Forst. Ins. 1844. 2 Ann. Soc. ent. France (2), iii. 1845, p. 355.
® Tr. ent. Soc. London, 1885, pp. 224, 219.
a mass of cocoons more or less loosely
of a stalked cocoon (Fig. 368) being
|
560 HYMENOPTERA CHAP. |
\
|
Gee PARASITICA 561
‘This family is of enormous extent; we have several hundred
‘species of it in Britain,’ and there are no doubt many thou-
sands of undescribed exotic forms. To Apanteles glomeratus. we
are indebted for keeping our cabbages and kindred vegetables
from destruction by the caterpillars of the white butterflies.
The larvae of the various species of Pieris, as well as those of
other Lepidoptera, are attacked by this little Insect, the masses of
whose cocoons may frequently be found in numbers in and near
cabbage gardens. The tropi-
eal species of Braconidae are
greatly neglected, but many
large and remarkable forms
—some of brilliant colours
—have been brought from
there, so that we are justified
in believing that Insects of 7.0
this family will prove to be |
very numerous. ‘There are
but few apterous Braconidae.
Both sexes of Chasmodon
apterus are destitute of wings;
the females of one species of
Spathius, and also those of (
Pambolus and Chasmodon are
apterous ; in a small number ,
of species of various genera
the wings are so minute as
to be incapable of serying as | |
organs of flight. In the |
genus Alloea the wings of
the male are shorter than
those of the female. |
Nae
Fic, 369.—Stenophasmus ruficeps, female.
Fam. VI. Stephanidae. Aru Islands.. (After Westwood.)
Antennae composed of many (thirty to seventy) joints; hind body
attached to the lower and posterior part of the median dorsal
1 A monograph of the British Braconidae was commenced by the Rey. T. A.
Marshall in 1885, and is still in progress, in. the Transactions of the Entomological
Society of London ; cf. op. cit. 1885, 1887, 1889, 1891, 1894.
VOL. V 20
ia
:
le ae a ee Pai ene. eee
5 62 HYMENOPTERA
‘|e
plate. Wings with a distinet costal cellule ; head globose,
posterior femora frequently toothed.
This is a doubtful family, consisting of a few onomalaaa
Insects. Schletterer assigns to it only two genera, Stephanus and ©
Stenophasmus;! both have a wide distribution over the world, —
though we have no species in Britain. Nothing is known of
their habits, and they are apparently all very scarce Insects.
The definition is compiled from those of Cameron and Schlet-_
terer. There seems Py. little to distinguish these Insects from
Braconidae. , :
Fam. VII. Megalyridae.
Hymenoptera with short broad hind body, which is not separated |
by a pedicel from the thorax. The female has a very long
bristle-like ovipositor. Antennae with fourteen jornts.
This family is constituted by the Australian genus Megalyra,?
one of the most interesting of the numerous extraordinary Insect-_
forms found in that region; the species appear to be very rare
and not numerous. Apparently nothing is known as to their
habits. It is quite possible that these Tnsects will prove to be
anomalous Braconidae.
ee ee ee
Fam, VIII. Evaniidae.
Petiole of the abdomen attached to the upper part of the median
dorsal plate; antennae not elbowed, of thirteen or fourteen
joints. Wings with a moderate number of nervures. Larva
of parasitic habits.
a a ee eS ee, ne id
This family is composed of only three genera—Zvania,
Gasteruption, and Aulacus, each possessing a considerable number
of species; they agree in the characters mentioned above, and
may be readily recognised by the peculiar insertion of the
hind body. This character occurs outside the limits of the
_Evaniidae only in one or two genera of Chaleididae and
Braconidae ; it is to this latter family that the Evaniidae must ;
be considered most closely allied. '
The species of the genus Evania are believed to live at
a)
a
j
1 Berlin entom. Zeitschr. xxxiii. 1889, p. 197. 2 Ibid. 7 |
CU PARASITICA 563
the expense of cockroaches (Blattidae), and to deposit their
eggs in the egg-capsules of those Insects. The species of
Gasteruption live, in the larval state, on the larvae of other
Hymenoptera, more especially of such as form nests in wood.
Very little is known as to the habits of the species of Aulacus,
but it is believed that they are parasitic on members of the
Hymenopterous families, Siricidae and Oryssidae. Only the
most meagre details as to the life history of any of the Evaniidae
have been recorded, The species of Evania are met with most
freely where cockroaches abound, and. are said, hence, to be
frequently observed on board ship. Two or three species of
each of the two genera Hvania- and Gasteruption occur in .
Britain. The latter genus is |
. more widely known under the |
name of Foenus.'
Fam. IX. Pelecinidae.
Sexes very different ; the female ) ‘ ’ ]
without exserted ovipositor, a a eh
but with extremely long ge rat | \
abdomen. Articulation — Bis |
between the femur and /
trochanter oblique and | :
elongate, but without divi-
sion of the trochanter.
|
This family at present com- |
prises, according to Schlet- )
terer,? only the three genera |
Pelecinus, Ophionellus, and
Monomachus. The systematic
position of the Insects is very
doubtful, and their habits are
pate rdithde Bown: ripest Fie, 370.—Pelecinus polyturator, q.
polyturator (Fig. 370) appears, patty
however, in the female sex, —
to be acommon Insect over a large part of the warmer regions
1 Monograph, Schletterer, Verh. zoo/.-bot. Ges. Wien, xxxv. 1885, p. 267, ete. ;
xxxvi. 1886, p. 1, etc. ; and Ann. Hofmus. Wien, iv. pp. 107, ete.
2 Berlin. entom. Zeitschr. xxxiii. 1889, p. 197.
564 HYMENOPTERA ¢ APL
of the New World; it is in all probability parasitic in its habits,
the elongate ovipositor of the female Ichneumon being in ie
Insect replaced by an extraordinary linear extension of the abdo- —
men itself. Doubleday has recorded that he saw twenty ¢
thirty specimens of this species that had perished with thei
elongated hind bodies inserted into the stem of a tree, fre nm I
which they could not extricate themselves. On the other hand, —
Patton thinks they are parasitic on locusts.’ hie 3
The male in Pelecinus has the proportions of the nani of
the body normal, there being no elongation of the abdomen; oa
thus differs very much in appearance from the female. Theiy
seems to be very little to distinguish Pelecinus from Procto- —
trypidae. The undivided trochanters have led to these Insects —
being placed, by some, among the Aculeate Hymenoptera, This"
character, as we have already shown, occurs ale in Procto- —
nee ,
Fam. X. Trigonalidae.
Abdomen ovate, not separated by a pedicel from the thorax.
Antennae twenty-five-jointed. Trochanters imperfectly two- —
jointed. Both the anterior and posterior wings provided with
a well-developed neuration. Abdomen composed of only five —
apparent segments. Larva (in some cases) parasitic on Aculeate —
Hymenoptera. a
fe
\
\
This family is chiefly constituted by the very rare Insects —
contained in the genus 7'rigonalys, of which we have one species —
in Britain. Although, so far as appear-
ance goes, they have little in common —
with the parasitic Hymenoptera, and _
look quite like members of the Acu- —
leata, yet the late F. Smith found a —
species in the cells of Polistes lanio, —
thereby showing it to be of para- —
sitic habits. Although some Aculeate —
Hymenoptera are also of parasitic —
Fic. 871.—Trigonalys maculi- habits, yet the characters of Trigonalys
[ron ah ee perhaps agree, on the whole, better —
with the Hymenoptera parasitica. The British species is very
Y Amer. Nat. xxviii. 1894, p. 895. See also Forbes, Rep. Ins. Illinois, xix. 1896, p. 79.
¥
XXIIT | PARASITICA 565
rare. The South American genus, Vomadina, looks still more like
a bee, and the trochanters are even more imperfectly divided
than they are in some of the Aculeate group, Nyssonides, the
outer portion being merely a small piece imperfectly separated
from the base of the femur.
Note——tThe citation of Saint Augustine on p. 85 is made in the words
used by Wasmann in Der Trichterwickler, eine naturwissenschaftliche Studie
diber den Thierinstinkt, 1884.
The authenticity of the passage we have adopted as the motto for this
volume is somewhat doubtful. It is explained in an “ admonitio ad lectorem ”
of the soliloquy, that this work is probably a compilation by a later writer,
from two, or more, works of Saint Augustine. Father Wasmann has been so |
kind as to inform the writer that the idea of the passage quoted occurs
frequently in the undoubted works of the Saint, as, for instance, de Civitate
Dei, lib. xi. cap. 22 ; Serm. ccxiii. in traditione symboli II. cap. i. ; contra
Faustum, lib. xxi. cap. v. etc. The passage quoted is, however, the only one
in which “angeli” and “vermiculi” are associated.
INDEX
Every reference is to the page: words in italics are names of genera or species ;
figures
in italics indicate that the reference relates to systematic position ; figures in thick
type refer to an illustration ; f. = and in following page or pages.
ABDOMEN, 109 ; of Hymenoptera, 492 f.
Abdominal appendages, 188, 189, 190
Acantherpestes, 74, 76, 80
Accessory glands, 392, 404
Acheta, 330, 338
Achorutes murorum, 194
Acini, 126
Acoustic orifice, 317
Acridiidae, 201, 279-310, 309
Acridiides, 310
Acridium peregrinum, 298 ; growth, 156 ;
at sea, 297—see also Schistocerca
Acrophyllides, 278
Aculeata, 520
Aculeates and Proctotrypids, 535, 564
Adler, on alternation of generations, 530 ;
on galls, 526 f. ; on useless males, 498
Aeschna cyanea, 412 ; A. grandis, labium,
411 ; nymph, 420, 421
Aeschninae, 416, 426
Agaonides, 547
Agathemera, 274, 276
Agrion nymph, 426
Agrion pulchellum, 412
Agrioninae, 412, 426
Agriotypides, 557
Agriotypus armatus, 557
Air sacs, 128, 282, 283, 294, 495
Alaptus excisus, 587; A. fusculus, 538
Alar organs, of earwigs, 206 ; of Blattidae,
225 ; of Mantidae, 245 ; of Phasmidae,
269 ; of Acridiidae, 281 ; development
of, in earwigs, 212; in Mantidae, 248
—see also Tegmina, Wings, Elytra
Alary muscles, 134
Albarda on Raphidiides, 448
Alder flies, 444
Aleuropteryx, 471
Alimentary canal, 123-127, 403, 446; of
may - flies, 438 f.; of Panorpa, 450;
closed, 457, 466, 496, 544
Alitrunk, 489 f., 490, 492
Alloea, 561
Alternation of generations, 497, 530
Amber, Myriapods in, 74, 76, 77 ; Insects,
179; Aptera, 196; Blattidae, 239;
Mantidae, 258; Phasmidae, 276; Pso-
cidae, 397 ; Perlidae, 407 ; Phryganeidae,
485 ; Tenthredinidae, 518; Cynipidae,
533.
Ambua, 40
Ameles, 245
Ametabola, 158, 174
-Amnion, 148, 545
Amnios, 291
Amorphoscelides, 251, 259
Amorphoscelis annulicornis, 251
Amphibiotica, 342
Amphientomum paradoxum, 397
Ampulex, abdomen, 492
Ampulla, 290
Amylispes, 76, 80
Anabolia furcata, mouth-parts, 475; A.
nervosa, larva, 476
Anabrus purpurascens, 321
Anaplecta azteca, folded wing, 227
Anaplectinae, 240
Anatomy — see External Structure and
Internal Anatomy
Anax formosus, 410, 414
Anderson, Dr. J., on Gongylus, 254
Anechura scabriuscula, 208
Anisolabis maritima, 205 ; A. moesta, 205,
A. tasmanica, 216
Anisomorpha pardalina, 274
Anisomorphides, 278
Anisopterides, 412, 414, 426; nymphs,
421
Anomalon, metamorphosis, 552
Anomalopteryx, 484
Anostostoma australasiae, 326
Anoura, 190
568
PERIPATUS-—MYRIAPODA——INSECTA
Ant, brain, 119; nervous system, 495 ;
castes, 500
Ant destroyer, 545, 559
Ante-clypeus, 93
Antennae, 97 ; growth of, 212
Anthophora retusc, parasites of, 544, 545
Anthophorabia retusa, 545
Ant-lions, 453, 454
Anurida ‘maritima, 194, 195
Aorta, 135, 134
Apanteles glomeratus, 561
Apatania, 481; A. arctica, A. muliebris,
481
Apex, 112
Aphilothriz, 531
Apocrita, 519
Apodeme, 103
Apophysis, 103, 520
Appendages, 91, 188, 189, 190
Aptera, 172, 180-189
Apterous Insects, 205, 216, 217, 220, 234,
235, 252, 261, 262, 264, 269, 272, 274,
277, 299, 302, 303, 307, 321, 322, 323,
324, 325, 326, 329, 518, 556, 561—see
also Wingless Insects
Apterygogenea, 175, 196
Aquatic Acridiidae, 301, 303; Aq. Hymen-
optera, 538, 557 ; Aq. Phasmids, 272
Arachnides antennistes, 77
Archidesmidae, 76
Archidesmus, 76
Archijulidae, 76
Archipolypoda, 74, 76
Arolium, 105, 223
Arrhenotoky, 141, 498
Arthromeres, 87
Arumatia ferula, anatomy, 262
Ascalaphides, 459 f.
Ascalaphus coccajus, 459 ; A. longicornis,
459 ; A. macaronius, 460 ; eggs, 460
Aschipasma catadronus, 263, 266
Aschipasmides, 278
Ashmead, on Mymarides, 537 ; on Procto-
trypids, 537 ; on Scleroderma, 536
Astroma, 300
Asymmetry, 216
Athalia (centifoliae) spinarum, 515
Atropinae, 394 f.
Atropos divinatoria, 394 f., 396
Atta (Oecodoma) cephalotes, 501
Attitude, 248, 250, 256, 268, 514
Attraction of light, 230
Auditory organ, 400; of Calotermes, 358
—see also Ear
Audouin on thorax, 100, 101
Aulacus, 562
Aulax, 532
Avicenna, 41
Axes of body, 113
BacILLIDES, 278
Bacillus patellifer, 263
“Ts
| Bittacus, 451, 453; B. tipularius, 452
Blaberides, 247
- Blood-gills, 479 @
Bacteria, 276
Bacteriides, 277
Bacunculides, 277
Baétis, 433
Ballostoma, 196
Ballowitz on spermatozoa, 140
Barber, Mrs., on 8. African locust, 294 —
Barbitistes yersini, 321
Barnston on Perlidae, 402, 405
Barriers with eggs, 461
Base, 112
Basement membrane, 162
Bassett on oviposition of inquilines, 532 *
Bataillon, on metamorphosis, 131, 168;
on reversed circulation, 135 4
Bates, on singing grasshopper, 319 ; on
Termites, 375 :
Bateson, on forceps of earwig, 209 ; on ;
antennae of same, 212 4
Batrachotettiz whiti, 305
Bedeguar, 527, 531
Bees killed by Locusta, 321 | :
Belt on domestic cockroaches, 231 ©§ |
Bermuda, 33
Bertkau, on Psocus, 391 ; on micropterous
Psocidae, 394
Bethylus habits, 5395
Bherwa, 326
Bird eaten by Mantis, 250
Bird-lice, 345, 351 a
Biting-lice, 345, 351
Blabera, 235; wings, 237; B. gigantea, iW
222 i
Black beetle, 221
Blanjulidae, 44
Blanjulus, 44
Blastoderm, 147
Blastophaga grossorum, 547 f.
Bilatta, 240 ;
Blattidae, 201, 220-241, 240; parasites
of, 563 ‘
Blattinae, 240 ; a
Blind Insects, 217, 233 ~
Blood, 132 é,
Blowfly, egg, 1453 roctainaraivaelel 163
Bolivar on eyes of Machilis, 185 id
Bombus, -dorsal vessel of, 133 ; metamor-
_ phosis, 497 ; B. ducorwm, 488
Bombyliidae, 291
Bonnet and Finot on Lugaster, 324
Book-lice, 390 f. ;
Boreus hiemalis, 451 ; larva, 453 ox
Boutan on concealment of leaf-like Tar “9
sects, 323
Brachyscelides, 526
Brachystola magna, 308
Brachytrypes megacephalus, 332
Bracon palpebrator, 559
Braconidae, 558 f.
INDEX
569
Bradford Cave, Myriapods in, 34
Brain, 118,120; of ant, 119; of Perlidae,
404
Branchiae, 401, 421—-see also Gills
Brandt on nervous system, 119, 495
Brauer, on classification, 175 ; on median
segment, 491 ; on hypermetamorphosis,
160 ; on Menorhyncha, etc., 161; on
Ascalaphus larva, 460; on development
of Mantispa, 464; on Palaeodictyop-
tera, 486; on Panorpa larva, 452; on
tegmina of Phyllium, 270
Breitenbach on Proscopiides, 299
Bridgman on British Ichneumonidae, 557 -
Brindley on growth of cockroach, 229
British, Myriapods, 36 ; Orthoptera, 201 ;
earwigs, 215; grasshoppers and locusts,
308; crickets, 339; Psocidae, 395;
Perlidae, 406 ; Odonata, 424 ; Sialidae,
444, 448 ; Chrysopides, 469 ; Trichop-
tera, 480 ; Phytophagous Hymenoptera,
504; Siricidae, 510; Cynipidae, 533 ;
Ichneumonidae, 557 ; Braconidae, 561
Brongniart, on fossil Insects, 428; on
fossil Neuroptera, 343;-on Neurop-
teroidea, 486; on post-embryonic de-
velopment of locust, 287; on young
Mantis, 247 f.
Brongniart and Becquerel on chlorophyll
in Phyllium, 268
Bruner on variation of Orthoptera, 804
Brunner, on Hypertely, 322 ; on classifica-
tion of Orthoptera, 202; of Blattidae,
240 ; of Mantidae, 259; of Phasmidae,
277; of Acridiidae, 309 ; of Locustidae,
328 ; on variation of Oedipoda, 304
Bryodema tuberculata, 281
Bugnion on histolysis, 166; on Hncyrtus,
545
Buller on Weta-punga, 326
Burchell on Mantis, 249
Burgess on. Psocus, 391 f.
Burmeister on Mantidae, 250
Bursa copulatrix, 139
CADDIS-FLIES, 473 f.
Caecum, 125
Caenis dimidiata, 442
Calcares, 104
Calepteryginae, 422, 426
Calepteryx, 417, 420, 422 ;
site, 538
Callimenides, 318, 329
Callimome bedeguaris, 532
Caloptenus spretus, 288 f., 289, 298, 303 ;
development, 289
Calotermes flavicollis, 362, 363, 371, 376 ;
C. nodulosus, 359; C. rugosus, 358,
382, 383
Calvert on Odonata, 412
Calvisia atrosignata, 266, 273 |
Calyx, 283, 439
its eggs’ para-
Cameron, on ant-parasite, 545; on gall-
producing plants, 527; on partheno-
genesis, 498, 499, 517; on Pimpla
larva, 557
Camerano on earwig, 211, 213
Campodea, 61; C. ert ylinus, 182, 183,
197
Campodeidae, 183
Camponotus, nerves, 495
Cannibalism, 425, 477
Cantharidae, 291
Capnia vernalis, 405
Caprification, 547 f.
Capsule of eggs, 201—see also Egg-capsule
Caraphractus cinctus, 538
Carboniferous, Myriapods, 75,76 ; Insects,
196, 238 f., 259, 276, 408, 428, 442 f.,
449
Cardiophorus larva, 90
Cardo, 95
Carnivorous and vegetarian, 250
Carpenter bee wings, 494
Carruthers on locust swarm, 292 .
Case, Hymenopterous, 514
Cases, caddis-fly, 476 f., 480, 481, 482,.
483, 484, 485
Castes, 500, 501
Caudal branchiae, 423
Cave, Myriapods, 34, 37; Insects, 197,
' 451; Locustidae, 321; cockroach, 232,
233
Cecidomyia, parasites of, 536, 537
Cenchri, 511
Centipedes, 30, 36, 40
Cephalocoema lineata, 299
Cephalonomia formiciformis, 536
Cephidae, 504 f.
Cephus integer, 505 ; C. pygmaeus, 505
Cerci, 110,183, 216, 257, 337, 400; of Blat-
tidae, 224, 238
Cermatia, 35
Ceroys saevissuma, 264
Cervical sclerites, 99, 99, 409
Chalcididae, 539
Chalicodoma muraria,
540 f,
Changing colour, 288, 253, 267, 268
Chasmodon apterus, 561
Chatin on labrum, 93 ; on mandibles, 95
Chauliodes, 447
Cheeks, 94
Cheimatobia brumata, parasites, 521
Chelidura dilatata, 205
Cheshire on fertilisation of bee, 499
Chilaspis lowii, 530; C. nitida, 531
Chilian Insects, 447, 463
Chilognatha, 30, 43, 47, 76 ; developmext
of, 63-72; structure of, 52-56, 53;
. double segments, 58, 70
Chilopoda, 30, 33, 44, 47, 52, 74, 74;
structure of, 56-59; development of,
70-72
nest, " parasites,
570 PERIPATUS——-MYRIAPODA—HINSECTA
Chitin, 162
Chitinogenous cells, 162
Chlorophyll in tegmina, 269
Choeradodis cancellata, 252
Cholodkovsky, on head, 87; on styles of
cockroach, 224; on embryolog y of
' Phyllodromia, 237; on morphology of
sting, 493
Chordeuma, 31
Chordeumidae, 44, 54
Chordeumoidea, 80
Chordotonal organs, 121
Chorion, 144
Chorisoneura, 240
Chromosomes, 146
Chrysopa eggs, 469 ; larva, 469; C. aspersa,
470; C. flava, 469 ; 0, pallida larva;.470
Chrysopides, 469 f., 472
Chun on rectal gills, 422
Chyle, 133
Chylific ventricle, 125, 228
Cimbex abdomen, 493 ; abdominal articu-
lation, 492; dorsal vessel, 134; C. syl-
varum, saws, 512
Cimbicides, 511, 517
Cinura, 182
Circulation, 132 f.; in caudal setae, 435
Cladomorphides, 278
Cladonotus humbertianus, 301
Classification, 171 f.; of Blattidae, 240;
of Mantidae, 259; of Phasmidae, 277 ;
of Acridiidae, 309 ; of Locustidae, 328 ;
of Gryllidae, 340
Claws, 105, 106, 469
Chitumnides, 278
Cloéon, eyes, 430; C. dimidiatum, larvule,
432; C. dipterum, nymph, 432 ; respira-
tion of nymph, 435
Clothilla, 395; C. pulsatoria, 395, 396 ;
anatomy, 392
Clypeus, 92, 93
Cockroaches, 220
Cocoons of sawfly, 515
Coeloblast, 149
Coleoptera, 173
Collembola, 782, 189 f.
Collophore, 193
Colobognatha, 44
Colour, 200
Commissures, 116
Common cocoons, 515
Compass Termite, 386
Complementary Termites, 361
Compound eyes, 97,430 ; (=facetted eyes)
in Myriapods, 36
Concealment by movement and position,
288 ; by selection of place, 308
Coniopterygides, 471
OConiopteryx lutea, 471;
471; OC. tineiformis, 472
Conocephalides, 313, 327, 328
Copiophora cornuta, 313
C. psociformis,
Cordulegaster, 415 ; C. annulatus, 415
Cordulegasterinae, 426
Corduliinae, 426
Correlative variation, 536
Corrodentia, 175, 389
Corrosion by Termites, 360
Corydalis, 447 ; C. crassicornis, 447
Corydaloides scudderi, 344
Corydia, 221; C. petiveriana, 233
Corydiides, 241
Coryna, 550
Corynothrix borealis, 191
Costa, 108
Cotes on Indian locusts, 298
Cotylosoma dipneusticum, 272
Coxa, 88, 104
Craspedosoina, 76
Crawlers, 447
Creepers, 407
Cretaceous Myriapods, 75; Insects, 485
Creutzberg on circulation, 436
Cricket, 330, 338
Crioceris asparagi, legs of larvae, 106
Crop, 114, 124, 495
Crunoecia irrorata, case of, 480
Cryptides, 557
Cryptocerus, abdomen of, 109
Cryptops, 36, 41
Crystalline cone, 98
Cuculligera flexuosa, 304
Cunningham on fig fertilisation, 549
Cursoria (Orthoptera), 201
Cuvier, 77
Cyclops form, 536
Cylindrodes campbellii, 336 ; C. kochi, 336
Cynipidae, 523
Cynips aciculata, 531; C. disticha, 530;
C. folii, 530; C. kollari, 530; ¢. lignt-
cola, 530 ; C. spongifica, 531
Cyphocrania aestuans, 266
Cyprus, 32
Cyrtophyllus concavus,320; C.crepitans,311
DaHt and Ockler on feet, 105
D’Albertis on may-flies, 441
Damsel-tlies, 417
Dancing may-flies, 439 f.
Dasyleptus lucasii, 196
Death-watch, 395 f.
Decaux on cannibalism of mole-cricket, 336
Deception, 250, 265
Decoys, 257
Decticides, 329
De Geer on earwigs, 214
Degeeriidae, 190
Deinacrida heteracantha, 326
Demoiselles, 417
Dendroleon pantherinus, 458
Denny on Mantis in England, 258
Derham on death-watches, 396, 397
Dermaptera, 202, 216
Dermatoptera, 202
‘
. ae
a
“ said
INDEX .
571
Derocalymma, 235
Deroplatys sarawaca, 248
De Saussure, on Orthoptera, 202 ; on wings
of Blattidae, 226 f.; on classification
of Gryllidae, 340 ; on Hemimerus, 217 ;
on nomenclature of Blattidae, 240; on
oceans as barriers to migration, 297
Desert Insects, 253, 304
Deuterotoky, 141, 497 f.
Deuto-cerebron, 118
Development, of alar organs of Platycleis,
312; of crickets, 332—see also Em-
‘bryology and Metamorphosis
Devonian, 428, 442
Dewitz on caste, 500; on ovipositor of
Locusta, 314; on morphology of sting,
493 ; on internal legs, 496 ; on develop-
ment of wings of Phryganeidae, 479,
480; on dragon-fly nymphs, 423; on
Chrysopa larva, 470
Diaphana fieberi, 226
Diapheromera femorata, 263, 264, 265, 267
Diastrophus, 532
_ Diaulus, 484
Dicranota, larva, glands of, 142
Dictyoneura, 277, 344
Dictyopteryz microcephala, 406; D. sig-
_ nata, 401
Dielocerus ellisii, 515
Digestion, 127
Dilarina, 465
Dilke, Sir Charles, on Orchis-like Mantis,
254
Dimorphic cocoons, 560 ; males, 547,549
Diplectrona, 479
Diploglossata, 217
Diplopoda, 43, 53, 74
Diploptera silpha folded wing, 227
Diptera, 172
Disgorgement, 495
Distant on §. African locust, 298
Ditrochous, 494, 520
Divided eyes, 409
Docophorus fuscicollis anatomy,
D. wterodes, D. cygni, 349
Dog, biting-louse of, 349 —
Dohrn on tracheal system of Gryllotalpa,
132; on embryology of Gryllotalpa,
336
Dolichopoda palpata, 322 °
Dorsal vessel, 133, 134; reversed action,
435
Dorsum, 100
Dragon-flies, 409
Drakes, 441
Drepanepteryx phalaenoides,
wings, 468
Drones, 499
Drummers, 237
Dubois on decapitated Mantis, 250
Duchamp on egg-capsule of cockroach, 228
Ductus ejaculatorius, 140
348 ;
453, 468 ;
Dudley and Beaumont on Termites, 372,
387
Dufour, on alimentary canal, 124; on tra-
cheal system, 129; on air sacs of Acri-
diidae, 283; on sexual organs, 138,
139; on testes, 140; on phonation,
286; on Tridactylus, 338; on Man-
tidae, 246; on earwigs, 210; on ana-
tomy of cockroach, 228 ; on anatomy of
Gryllotalpa, 335; on anatomy of Ter-
mites, 360; on anatomy of Panorpa,
450 ; on larva of Sialis, 446 ; on Myrme-
leon larva, 458
Duns, 441
Dust-lice, 390 f.
Dwellings of Termites, 385 f.
Dytiseus, mesothorax, 101 ; egg-tube, 138,
199)
Dzierzon theory, 499
Ear, 101, 121 ; of Acridiidae, 285 f., 285 ;
of Locustidae, 316 f., 316, 317; of
crickets, 332 ; of Gryllvtalpa, 333, 334
Earliest pre 238
Earwig, 202 f., 211, 213,.214; forceps,
208 f., 209 ; wing, 206; the name, 214
Eaton, on nymph, 157 ; on Ephemeridae,
435, 437, 440
Ecdysis, 156, 162; nature of, 169
Ectobia, 236 ; E. lapponica, egg-capsule, 229
Kctobiides, 240
Ectoblast, 149
Ectoderm, 148 ; of Peripatus, 20 f., 22
Ectognathi, 189
Ectotrophi, 189
Eggs, 143-145 ; of Peripatus, 19 ; of Myria-
pods, 38, 39, 64; of Ascalaphus, 460;
growing, 513; of parasites, 552 ; of egg-
parasites, 545 ; of Corydalis, 447 ; of Cyni-
pidae, 528 ; of Limacodes, 153; of Mallo-
phaga, 348 ; of Microcentrum, 314; of Phas-
midae, 265, 270 f., 270; of Perla, 404;
of Sialis, 445 ; of Trichoptera, 476 —
Egg-capsule, 265, 290 ; of Phylliwm, histo-
logy, 271
Egg-parasites, 522, 586, 538
Egg-tubes, 137, 139, 392—see also Ovaries
Eileticus, 76, 80
Eisig on chitinous excretion, 180, 163
Ejaculatory duct, 392, 414
Ejection of fluid, 264, 324, 399, 515
Elasmosoma, 559
Elater larva, 29
Elipsocus brevistylus, 393
Elytra, 108
Embia, 352, 353
Embidopsocus, 395
Embiidae, 351, 395
Embryology, 145-153 ; of Peripatus, 19 f.;
of Myriapods, 63 f. ; of parasites, 522:
of earwig, 216; of Blattidae, 237 ; of
Encyrtus, 546 ; of Gryllotalpa, 336 ; of
572
PERIPATUS—-MYRIAPODA—-INSECTA
Polynema, 588; of Smicra, 545; of
Proctotrypidae, 536 f.
Emergence from egg, 263, 264, 290,291, 313
Empodium, 105
Empusa pauperata, 245, 257
Empusides, 259
Encyrtus fuscicollis development, B45
Endoblast, 149
Endoderm, 148; of Peripatus, 20 f., 22
Endolabium, 97
Endo-skeleton, 399
Eneopterides, 340
Enock on Alaptus and Cansohemeds 538
Enoicyla pusilla, 481
Entognathi, 189
Entomology, 86
Entothorax, 103, 114, 116
Entotrophi, 189
Eocene, 407
Eoperipatus, 24 n.
Eoscolopendridae, 80
Ephemera, 434; H. danica, 429, 441;
wing, 431; #. vulgata, 441; nymph, 433
Ephemeridae, 429-443
Ephippigera Malpighian tubes, 335; £.
rugosicollis, 323
Ephippigerides, 318, 329
Epiblast, 65, 149
Epicranium, 92, 93, 93 ©
Epidemes, 107
Epilamprides, 240
Epimeron, 100, 101, 104
Episternum, 88, 100, 101, 104
Epistome, 92
Epithelium of stomach, 126
Eremiaphila, 2438, 253 ; E. vs bien 253
Eremobiens, 304
Erianthus, 301
Erichson on Neuroptera, 342
Erne on Rhyssa, 554
Etoblattina manebachensis, 238, 239
Eucharis myrmeciae, 545
EKuchroma, head and neck 99
Eucorybas, 37
Eugaster guyoni, 324
Eugereon bockingi, 486
Eumegalodon blanchurdi, 327
Eumegalodonidae, 327
Euorthoptera, 216
Euphaea, 422
Euphoberia, 76, 80
Euphoberiidae, 73, 76
Euprepocnemis plorans, 303
Eurycantha australis, 274
Eurypauropidae, 47
EKurytoma abrotani, 539
Eusthenia spectabilis, 407 .
Eutermes, 374 ; EL. ripperti, 388
Euthyrhapha, 296
Evania, 562
Evaniidae, 562
Exner on sight, 416
Exodus, locust of the book of, 298 ©
Exsertile blood-saes, 132
External parasite, 555
External structure, 87; diagram, 88; of
earwigs, 203 f. ; of cockroaches, 221 ; :
‘of Mantidae, 242 £3) ef Phasmidae,
260 f. ; of Acridiidae, 280 f.; of Odon-
ata, 409 f. ; of Ephemeridae, 430 f. ; of
Panorpa, 450; of Phryganeidae, 474 ;
of Hymenoptera, 489 f.; of Tenthre-_
dinidae, 511 é
Eyes, 97—see also Compound Eyes and aa,
Ocelli ;
FABRE on Leucospis, 540; on Monodon-
tomerus, 543; on Sirex, 509 :
Facetted eyes—see Compound Eyes:
Family, 177
Fasting, 448, 458
Fat-body, 136
- Feeding, by Termites, 376 ; young, 495
Femur, 88, 104
Fenestra, 221
Fenestrate membrane, of eye, 98 ; of aie
cardium, 134
Fertilisation, 499 ; of fig, 549
' Field- cricket, 332
Fields of wings, 206
Fig-Insects, 547 f.
. Figitides, 525
- Finot on Japyx, 196
Fire-brats, 186
. Fischer on instars, 158
Fish destroyed, 425
Fletcher on parthenogenesis, 498
Flight, 416
Floral simulators, 254 f.
Flying-machine, model for, 417
Foenus, 563
Foetus of Hemimerus, 218
Foramen, occipital, 92, 94.
Forbes on Blattid, 235
Forceps of earwigs, 208, 209 . .
Forel on nervous system of ant, 495
Forficula auricularia, 202 f., 204, 209,
211; F. gigantea, 210
Forficulidae, 201, 202
Formica-leo, 456
Formicajo, 456
Formicario, 456
Fossil, Insects, 178, 472, 485, 486 ; Acri-
diidae, 308 ; Blattidae, 238 ; : cricket,
340; dragon-flies, 427; earwigs, 216 :
Locustidae, 328 ; Mantidae, 258 ; may-
flies, 442, 443; Phasmidae,. 276;
Panorpidae, 453; Perlidae, 407 ; Siali-
dae, 449; Termites, 389; Thysanura,
196 ; Myriapods, 72 £.; Palaeozoic Neu-
roptera, 343
Founding communities, 381
Fourmilions, 456
' Fowl, biting-louse of, 350
INDEX
573,
Fritze on Ephemerid alimentary canal, 439
Frons, 94
Front wings absent, 260 f.
Fungus chambers, 387
Fungus-growing Termites, 885, 387
Funiculus, 492
Furea, 103
Fureal orifices, 399, 402
GALAPAGOS Islands, 459
Galea, 95
Gall- flies, 523 f.
Galls, 514 f. ; nature of, 525 f., 533
Ganglia, 116.
anin, on metamorphosis, 162 ; on embry-
ology, 536 f., 538
Gasteruption, 562
Gena, 94
Geophilidae, 46, 58, 7%
Geophilus, 33, 36, 39, 46; marine, 30 ;
phosphorescent, 34
Geoscapheusides, 241
Gerascutigeridae, SO
Gerephemera simplex, 428
Gerstaecker, on Neuroptera,
mouth of Odonata, 411
Giebel on Mallophagea, 347
Gigantic Insects, 276, 306, 428
Gilbert White, on mole-cricket, 333; on
field-cricket, 339
Gills, 132, 400, 421, 432f., 478; jointed, 445,
446, 467; filamentous, 476; spongy,
447 ; prothoracic, 443; of pupa, 483 ;
on imago, 401, 479 ; blood-gills, 479
Giraud on Cynipid oviposition, 528
Gizzard, 124, 125
Glacier water, 405
Glande sébitiqne, 139
Glands, 139, 142 ; conglobate, 229 ; maxil-
lary, 458; mushroom, 258—see also
Salivary Glands
Glandulae odoriferae, 31, 36, 54
Glomeridae, 43, 76, 8O
Glomeridesmidae, SO ~
Glomeris, 33, 43, 52
Gnathites, 94, 97
Golden-eyes, 469
Géldi on eggs of Phasmidae, 265
Gomphinae, 426
Gomphocerus, 308
Gomphus, 415
Gonapophysis, 110
Gongylus gongyloides, 254 f., 255
Gosch on median. segment, 491
Goureau on Microgaster, 560
Graber, on dorsal vessel, 134; on blood
cells; 137; on embryology, 148-151 ;
on ears, 286; on ears of Locustidae,
316, 317; on chordotonal organs, 121 ;
on blood, 133; on phonation of Steno-.
bothrus, 284 ; on Platycleis, 312
Grassi, on Myriapoda, 47 ; on Campodea,
343; on
163; on Hinbia, 353 ;
361 f.
Grassi and Rovelli on Thysanura, 182
Green grasshoppers, 311
Green, Mr. Staniforth, on
larva, 461 ;
Gromphadorhina portentosa, 235
Grosse on Mallophaga,. 346
Growth of wings, 393; of Mantidae, 248
on Termitidae,
Helicomitus
-Gryllacrides, 329
Gryllidae, 201, 330-340, 340
Gryllides, 340
Gryllotalpa, 332; dorsal vessel, 134 ;
Malpighian tubes, 127 ; tracheal system,
132
Gryllotalpides, 340
Gryllus, head, 93; G. campestris, 332,
339 ; G. domesticus, 330, 338
Guilding on Ulula, 461
Gula, 88, 93 4
Gyri cerebrales, 119
Gyropus, 350
HAASE on abdominal appendages,
192
Haemocoele, 22, 23
Hagen, on segments, 88; on wing-rudi-
ments, 395 ; on respiration of immature
dragon-fly, 423 f. ; on larvae of Ascala-
phides, 460 ; on amber Psocidae, 397 ;
on Platephemera, 428; on Perlidae,
401; on Psocidae, 393 f. ; on Termites,
360 f.
Haldmanella, 308
Halesus guttatipennis, 473
Haliday on Bethylus, 535
ITalobates, 83
Halteres, 108
Iiansen on Hemimerus, 217
Haplogenius, 461
Haplophlebium, 345
Haplopus grayi, egg, 265
Harpagides, 259
Harpalus caliginosus, head, 92
Harpax ocellata, 253; H. varieyatus, 244
Harrington on Oryssus, 507
Harris on Katydids’ music, 320
Hart on forms of Atta, 501
Hartig on gall-flies, 530
Harvesting Termites, 383
Harvey on metamorphosis, 168
Hatchett Jackson on ecdysis, 162; on
oviduct of ee tae 139
Haustellata, 94
189,
Haustellum, 476
Haviland on Termites, 368, 373, 384
Hawaiian Islands, 354, 395, 425, 471
Head, 92-94
Heart, 133
Heat, 131
Helicomitus insimulans, 460, 461
Helicopsyche shuttleworthi, cases of, 482
574
PERIPATUS—MYRIAPODA——-INSECTA
Hellgrammites, 447
Helminthomorpha, SO
Helorus anomalipes, 534
Hemerobiidae, 453 f.
Hemerobiides, 465 f,
Hemerobiina, 467, 472
Hemerobius larva, 467
Hemichroa rufa, 498
Hemimeridae, 207, 217
Hemimerus hanseni, 217 ; foetus of, 218 ;
H. talpoides, 218
Hemimetabola, 158
Hemiptera, 173
' Hemiteles, 556
Henking on embryology, 145
Henneguy on egg-capsule of Phyllium,
271 ; on embryology of Smicra, 545
Heptagenia, 440; H. longicauda, 437
Hessian-fly, parasites, 537
Heterogamia, 222; H. aegyptiaca, 220;
egg-capsule, 229
Heterometabola, 158
Heteromorpha, 158
Heterophlebia dislocata, 427
Heteropteryx grayi, 262
Hetrodides, 329
Hexapoda, 86
Heymons on earwig embryology, 216
Hind body, 109
Hind wings absent, 429
Histoblasts, 167
Histogenesis, 165
Histolysis, 165, 166 -
Hodotermes japonicus, 383;
landi, 384; H. mossambicus, 356 ;
brunneicornis, 359 ;
371
Hoffbauer on elytra, 108
Holocompsa, 226, 235
Holometabola, 158
Holophthalmi, 459
Homomorpha, 158
Hooks for wings, 494
Hoplolopha, 303
Hose, 393
Howard, on pupation of Chalcididae, 550 ;
on Hydropsyche, 483
Hubbard and Hagen on Termites, 388
Humboldt, 31
Humpback, 445
Huxley, on head, 87 ;
99
H. havi-
H.
on cervical sclerites,
Hydropsyche, 479
Hydropsychides, 482 ;
Hydroptila an gustella, 474 ;
lant, larva, 484
Hydroptilides, 484
Hylotoma rosae, 513
Hymenoptera, 173, 487-565
Hymenoptera phytophaga, 503 f.
Hymenopus bicornis, 253
Hyperetes, 395, 397
larva, 483 ‘
H, maclach-
H, quadricollis, -
Hypermetamorphosis, 158, 159, 465, 540,
552, 557
Hyperparasitism, 521
Hypertely, 323
Hypnorna amoena, 234
Hypoblast, 65, 149
Hypocephalus, 99
-Hypochrysa, 470
Hypodermis, 162, 480
Hypoglottis, 96
Hyponomeuta cognatella, parasite of, 545 |
Hypopharynx, 96—see also Lingua
ICHNEUMONES ADSCITI, 559
Ichneumon-flies, 265, 531; uninjurious
264 ; supplementary, 558
Ichneumonidae, 551-558
Ichneumonides, 557
‘| Ictinus, 419
Ilyodes,. 80
Imaginal, dises, 165, 166 ; folds, 165
Imago, 157
| Imbrications 493
Imhof on Perla, 403 f.
Inaequipalpia, 480
Indusial limestone, 485
Infra-oesophageal ganglion, 11
Inner margin of wing, 108
Inocellia, 447
Inquilines, 373, 524, 531, 533
Insecta, definition, 86
Instar, 155, 158
Instinct of Leucospis, 541
Integument, 162
Internal anatomy, 186 f. ; of f Acridiidae,
282 f.; of earwigs, 210; of Gryllotalpa,
335 ; of Hymenoptera, 494 ; of Libel-
lula, 414; of Mantidae, 246; of Myr-
meleon larva, 457, 458; of Odonata,
~ 414; of Stylopyga orientalis, 228 ; of
Phasmidae, 262; of Raphidia, 448 ; of
Sialis larva, 446 ; of Thysanura, 187 f.
Intestine, 114, 124
Involucrum alarum, 206
Iris oratoria, 248
| Isogenus nubecula, 405, 406
| Isopteryx, 400
Isosoma, 546
Isotoma, 190
JAMAICA, 388
Japygidae, 184
Japyz,abdomen of,109; J.solifugus, 184,196
Jhering, Von, on Termites, 387 .
Joint, 105
Joint-worms, 546
Joly on Ephemeridae, 431; on anatomy
* of Phyllium, 262
Julidae, 34, 43, 71, 73, 77
Juloidea, 80
Julopsis, 74 ;
Julus, 36-39, 52; J. nemorensis, 43; J.
terrestris, 37, 70, 773; breeding, 37;
}
INDEX —
575
development, 66-69 ; heart, 50; ovum,
63, 64; eye, 69
Jurassic, 216, 259, 407, 442
Jurine on pieces at base of wing, 102
KAMPECARIS, 76
Karabidion, 274
Katydids, 319, 320
King, 361, 378
Klapalek, on Trichopterous larvae, 484 f. ;
on Agriotypus, 557.
Knee, 104
Koch, 42
Koestler on stomatogastric nerves, 120
Kolbe, on entothorax, 103 ; on wings of
Psocidae, 394
Kollar on Sirex, 509
Korotneff on embryology of Gryllotalpa,
336
Korschelt on egg-tubes, 138
Korschelt and Heider on regenerative
tissue, 167
Kowalevsky, on phagocytes, 166; on re-
generative tissue, 167 ; on bee embryo,
496
Kradibia cowani, 549
Krancher on stigmata, 111
Krawkow on chitin, 162
Kulagin, on embryology, 537 ; of Encyrtus,
545
Kiinckel d’Herculais, on histoblasts, 167 ;
on emergence of Stawronotus, 290
Labia minor, 214
Labidura riparia, 210, 211, 214, 215
Labium, 95; of Odonata, 410, 411; of
O. larva, 420
Laboulbéne, on Anurida maritima, 194 ;
on Perla, 399
Labrum, 93, 93
Lacewing flies, 453, 469
Lachesilla, 395
Lacinia, 95
Laemobothrium, 347
Lamarck, 77
Lamina, subgenitalis, 224 ; supra-analis, 224
Landois on stigmata, 111 —
Languette, 96
Lankester, 40
Larva, 157 ; (resting-larva), 164 ; oldest, 449
Larvule, 431, 432
Latreille, 30
Latreille’s segment, 491
Latzel, 42, 77
Latzelia, 80
Leach, 30, 77
Lead, eating, 510
Leaf-Insects, 260
Legs, 104 ; internal, 496 ; four only, 549;
of larvae, 106, 110
Lendenfeld, on dragon-flies, 416, 417 ; on
muscles of dragon-fly, 115
Lens, 98
Lepidoptera, 173
Lepisma, 185, 196; L. saccharina, 186; L.
niveo-fasciata, 195
Lepismidae, 185
Leptocerides, 482
Leptophlebia cupida; 430
Lespés on Calotermes, 364°
Leuckart on micropyle apparatus, 145
Leucocytes, 137
Leucospis gigas, 540;
habits, 540 f.
Lewis, Geo., on luminous may-fly, 442
Lewis on Perga, 518
Leydig, on brain, 119, 120 ; on Malpighian
tubes of Gryllotalpa, 335 ; on ovaries,
137, 142 ; on glands, 142
Lias, 216, 236, 340, 427, 428, 453, 485, 503
Libellago caligata, 413
Libellula quadrimaculata, 411, 425
Libellulidae, 409
Libellulinae, 416, 426
Lichens, resemblance to, 253
Liénard on oesophageal ring, 118
Light, attraction of, 441
Ligula, 96
Lilies and dragon-flies, 426
Limacodes egg, 153
Limacomorpha, 80
Limnophilides, 487
Lingua, 95, 96, 391, 411, 420, 437
Linnaeus quoted, 84
Liotheides, 346, 350
Lipeurus heterographus, 346 ; L. bacillus,
347; L. ternatus, 349
Lipura burmeisteri, 190 ; L. maritima, 194
Lipuridae, 190
Liquid emitted, 264, 324, 399, 515
Lissonota setosa, 551
Lithobiidae, 45, 70, 75
Lithobius, 32, 36-39, 41, 45, 58; breeding,
38 ; structure, 48, 49, 57
Lithomantis, 259 ; L. carbonaria, 344
Locusta, ovipositor, development and struc-
ture, 315; L. viridissima, 318, 319, 321,
324, 327
Locustidae, 201, 311-329, 328
Locustides, 329
Locusts, 291 f.; of the Bible, 298; in
England, 299; swarms, 292-299 ; eggs,
292
Loew on anatomy of Panorpa, 450 ; of
Raphidia, 448
Lonchodes duivenbodi, ege, 265; L. nema-
todes, 260, 261.
Lonchodides, 277
Longevity, 377, 429, 438; of cockroach,
229
Lopaphus cocophagus, 264
Lophyrus pini, 511
Low on Coniopteryz, 471, 472
Low, F., on snow Insects, 194
larva, egg, 542 ;
576
PERIPATUS—MYRIAPODA—INSECTA
Lowne, on embryonic segments, 151; on
integument, 162; on stigmata, 111; on
respiration, 130
Lubbock, Sir John; on Pawropus, 62; on
aquatic Hymenoptera, 538 ; on auditory
organs, 121; on sense organs, 123 ; on
respiration, 180; on stadia, 1653; on
Cloéon, 432, 437 ; on Collembola, 192 ;
on Insect intelligence, 487
Lucas on mouth-parts of Trichoptera, 475
Luminous may-flies, 442
Lycaenidae, eggs, 144
Lyonnet on muscles, 115
Lysiopetalidae, 76
MACHILIDAE, 184
Machilis maritima, 185 ; M, polypoda, 184
Macronema, 478
Malacopoda, 77
Mallophaga, 342, 245-350
Malpighi on galls, 525
Malpighian tubes, 114, 124, 127, 187,
358, 360, 392, 403, 414, 421, 448, 457, ©
458; of Gryllotalpa, 335; of Ephippigera,
335; of Mantis, 246; of Myriapods,
48
Malta, Myriapods at, 35
Mandibles, 94, 95; absent, 474, 475
Mandibulata, 94
Manticora, 304
Maatidae, 201, 242-259, 259
Mantides, 259
Mantis, immature tegmina, 248 ; parasite,
546; M. religiosa, 246, 247, 258
Mantispa areolaris, 463; M. styriaca larva,
464
Mantispides, 463 f.
Mantoida luteola, 251
Marchal on Malpighian tubes, 127
Marine Myriapods, 30
Marshall, on Apanteles cocoons, 560; on
Braconidae, 561
Mask, 420
Mastacides, 301, 309
Mastax guttatus, 301
Maternal care, 214, 336, 517
Maxilla, 95, 96 ; of Odonata, 411 ;
190
May-flies, 429 ; number of, 442
Mayer, on Apterygogenea, 196 ;
fication, 547, 548
Mazon Creek, Myriapods at, 75
M‘Coy on variation of ocelli, 267
M‘Lachlan, on Ascalaphides, 459; on
Oligotoma, 354; on Psocidae, 395; on
Trichoptera, 480 f.
Mecaptera, 174, 453
Mechanism of flight, 416
Mecistogaster, 412
Meconema varium, 321
Meconemides, 328
Mecopoda, 319
absent,
on capri-
Mecopodides, 328
Mecostethus grossus, 285, 299, 308
Median plate, 504, 506, 507, 512
Median segment, 109, 490, 491
Megachile, nervous system, 496
Megaloblatta rufipes, 235
Megalomus hirtus, 468
Megalyra, 562
Megalyridae, 562
Meganeura monyi, 428
Megasecopterides, 344
Megastigmus, 547
Meinert, on earwigs, 210, 211, 212; on
Myrmeleon larva, 457 ; on stink-glands,
210
Melittobia, 545
Melliss on Termite of St. Helena, 389
Melnikow on eggs of Mallophaga, 348
Membranule, 413
Menognatha, 161
Menopon leucostomum, 348 ; M. pallidum,
350
‘Menorhyncha, 161
Mentum, 95, 96, 96
Mesoblast, 20, 65, 149
Mesoderm, 20, 149
Mesonotum, 88
_ Mesopsocus unipunctatus, I94
Mesothoracic spiracle, 491
Mesothorax, 101
‘Mesozoic, 309, 449, 485
Metabola, 158, 174
Metagnatha, 161
Metamorphosis, 153-170 ; of Hymenoptera,
497 ; of nervous system, 495 f.
Metanotum, 88
Metapodeon, 491
Methone, 200; M. anderssoni, 305, 206
Miall, on imaginal discs, 165, 167 ;
unicellular glands, 142
Miall and Denny, on pericardial tissue,
135; on epithelium of stomach, 126;
on spermatheca of cockroach, 228; on
stigmata, 111 ; on stomato-gastric nerves,
120
-Miamia bronsoni, 449
Microcentrum retinerve, 313, 314, 320
Microgaster, 559; M. fulvipes, 560; M.
globatus, 560
Micropterism, 339, 394, 405 f., 484
Micropyle, 145 ; apparatus, 404
Migration, 293, 425
Migratory locusts, 292, 297
Millepieds, 41
Millipedes, 30, 40, 41
Miocene, 216, 258, 407
Molanna angustata, mandibles of pupa,
477
Mole-cricket, 333 ; leg, 333
Moniez on Anurida maritima, 194
Monodontomerus, 532; M. cupreus, 543;
M. nitidus, 544
‘
ee A eT ee ee ee &.
HT
INDEX
577
Monomachus, 563
Monomorphic ant, 498
Monotrochous trochanters, 494, 520, 564,
565
Mordella eye, 98
Mormolucoides articulatus, 449
Morton, on gills of Trichoptera, 483 ; on
Perlidae, 406
Moult, 156
Moulting, 437 ; of external parasite, 556
Mouth-parts, of dragon-fly, 411 ; of dragon-
fly nymph, 420; atrophied, 430 .
Miiller, Fritz, on caddis-flies, 482 f.; on
fig-Insects, 549 ; on Termites, 358, 360,
374, 381, 382
Miiller, J., on anatomy of Phasmidae, 262
Murray, on Phyllium scythe, 263; on
post-embryonic development of Orthop-
tera, 265
Musca, metamorphosis, 163, 167
Muscles, 115
Music, of Locusta, 318; of Tanana, 319;
of Katydids, 319—see also Phonation
Mylacridae, 239
Mymarides, 537, 538
Myoblast, 149
Myriapoda, 27, 42, 74; definition, 29; as
food, 31 ; habits, distribution, and breed-
ing, 29-40 ; locomotion, 40 ; names for,
41 ; classification, 42-47 ; structure, 47-
63; embryology, 63-72; fossil, 72-77 ;
affinities, 78
Myrmecoleon, 456
Myrmecophana fallax, 323
Myrmecophilides, 340
Myrmeleo, 456
Myrmeleon, 456; M. europaeus, 457 ; M.
Sormicarius, 455, 457; M. nostras, 457 ;
M. pallidipennis, 456
Myrmeleonides, 454 f.
Nasutt, 370
Necrophilus arenarius, 462
Necroscides, 278
Needham on locusts at sea, 297
Nematus, 514; .V. curtispina, 498
Nemobius sylvestris, 339
Nemoptera ledereri, 462; NV. larva, 462
Nemopterides, 462
Nemoura, 401; N. glacialis, 405
Neoteinic Termites, 362, 380
Nervous system, 116
Nervures, 107, 108, 206 ; of Psocidae, 393 ;
of Embiidae, 352 ; of Termitidae, 359
Neuroptera, 172, 341-485; N. amphibio-
tica, 342; N. planipennia, 342
Neuropteroidea, 486
Neuroterus lenticularis, 523
Neuters, 137
Newman on abdomen, 491
Newport on |
of nymphs of Odonata, ©
LD ye ate ae ON a
INDEX
581
centrum, 313 ; on ovipositing of locust,
290 ; on subimago, 437; on Thalessa,
554
Ritsema on Hnoicyla, 481
Ronalds on anglers’ flies, 441
Roux on Necrophilus, 462
Royal pairs, 377 :
Riihl on earwig, 213
Sacs—see Air Sacs
Sagides, 328
Salivary glands, 124, 126, 187, 210, 228,
246, 283, 335, 348, 353, 403, 414, 495 ;
of Peripatus, oa s0f ‘Myriapods, 48,
49
Salivary receptacle, or reservoir, 126, 228,
246, 335, 348, 360
Saltatoria (Orthoptera), 201
Sandwich Islands—see Hawaiian Islands
Saunders, Sir Sydney, on. Scleroderma,
536 ; on caprification, 548
Saussure, H. de—see De Saussure
Savage on Termites, 368
Saw, 493, 512
Sawflies—see Tenthredinidae
Scales, 185, 189, 397
Scapteriscus, 334
Scelimena, 301
Schindler on Malpighian tubes,
Gryllotalpa, 335
Schistocerca peregrina, 298 ; development,
287 ; S. americana, 298, 308
Schizodactylus monstrosus, 325
Schizophthalmi, 459
Schizotarsia, 35, 46, 57, 58, 70,75 ;
ture, 59
Schletterer on parasitic Hymenoptera, 562,
563
Sclerite, 91
Scleroderma, 536
Scolia, ovaries, 138
Scolopendra, 30, 31, 32, 41, 78
Scolopendrella, 47, 61
Scolopendrellidae, 33, 42, 46
Scolopendridae, 31, 33, 39, 45, 75;
matophores, 39
Scorpion-flies, 449 f.
Seudder, on grasshopper music, 287 ; on
Katydids’ music, 320; on locusts at
sea, 297 ; on reproduction of lost limbs,
265; on fossil Insects, 486; on fossil
earwigs, 216 ; on fossil may-flies, 443 ;
on fossil Sialidae, 449; on Tertiary
Insects, 179 .
Scutellum, 100, 101
Scutigera, 35, 36, 48 ; sense organ, 51
Seutigeridae, 35, 36, 40, 46, 50
Scutum, 100, 101
Secondary, 427, 472 ;
Securifera, 503
Segmentation,
Smicra, 545
246; of
struc-
sper-
larva, 542
149, 237; of ovum of
Segments, 88, 90 ; number of, 87
Selys, De, on dragon-flies, 425, 427
Semi-pupa, 497
Sense organs, 121-123
Senses, 541, 544, 553
Sericostomatides, 474, 482
Series, 177, 201
Serosa, 148.
Serrifera, 503
Sessile abdomen, 493
Sessiliventres, 492, 496, 503
Sex, 498, 499, 500
Sexes, 137
Sexual organs, external, 141
Shaw on Orthoptera, 201
Sialidae, 407, 444
Sialides, 444
Sialis lutaria, 444 ; eggs, 445, larva, 445 ,
tracheal gill, 446
Silk, 127
Silo, parasite of, 558
Silurian Insect, 238
Silver fish, 186
Simple eyes, 97, 184—-see also Ocelli
Sinel, on marine Geophilus, 30
Siphonaptera, 174, 175
Sirex, habits of its parasite, 554; S. augur,
509; S. gigas, 508, 510; S. juvencus,
508
Siricidae, 507; parasites of, 563
Siricides, 510
Sisyra, 467 ; S. fuscata larva, 467
Sisyrina, 467
Sitaris humeralis, early stages, 159
Sloane, Sir Hans, on locusts at sea, 297,
Smallest Insect, 537
Smeathman on Termites, 366 f., 381, 383,
387
Smicra clavipes embryology, 545,
Smith, F., on Cynips, 530; on Trigonalys,
564
Smynthuridae, 797
Smynthurus. variegatus, 19155 S. fuscus,
192
Snow-Insects, 194
Social, Insects, 85, 361, 369 ; Gb TIeT eS:
488, 500 f.
Soldiers, 370, 371, 372
Somites, 87
Sommer on Macrotoma, 163, 195
Soothsayers, 242
Sound production, 358—see also Phona-.
tion
Spathius, 561
Species, number of—see Number
Spencer, Herbert, on caste and sex, 500
Spermatheca, 139, 228, 499
Spermatophores, 10, 39
Spermatozoa, 140
Sphaeropsocus kunowii, 397
Sphaerotherium, 43
Sphex chrysis, 490
582
PERIPATUS—MYRIAPODA—INSECTA
Spiders eaten, 464, 465
Spinneret, 458
Spinners, 441
Spiracles, 89, 111, 128 ; number of, 186 ;
of dragon-fly nymph, 423: absent, 436
—see also Stigmata
Spiral fibre, 128
Spongilla fluviatilis, larva in, 467
Spontaneous generation, 525
Spring of Collembola, 191
Spurs, 104
Stadium, 155, 158
Stalked, cocoons, 560; eggs, 469
St. Augustine quoted, 85, 565
Stein on Raphidia larva, 448
Stelis, parasitic, 544 ; parasitised, 543
Stem sawflies, 504
Stenobothrus, 308 ;
284
Stenodictyopterides, 344
Stenopelmatides, 321, 329
Stenophasmus ruficeps, 561
Stenophylla cornigera, 257, 258
Stephanidae, 567
Stephanus, 562
Sternum, 91, 100
St. Helena, 389
Stick-Insects, 260
Stigma of wing, 524, 534
Stigmata, 88, 89, 111, 204 ; position, 493 ;
on head, 193 ; 8. repugnatoria, 36—see
also Spiracles
Sting, 493 ; and ovipositor, 534
Stink-flies, 469
Stink-glands, 31, 125, 210, 264, 335
Stipes, 95
Stoll on spectres, etc., 254
Stomach, 114, 124, 125
Stomato-gastric nerves, 120, 121
Stomodaeum, 123, 151
Stone-flies, 407
Stratiomys strigosa parasite, 545
Stridulation, 304—see also Phonation
Stummer-Traunfels on Thysanura and
Collembola, 189
St. Vincent, island of, 461
Styles, 224, 238
Stylopyga ortentalis, 223, 228, 231, 236
Sub-imago, 429, 437 ;
Sub-Order, 177
Subulicornia, 426
Sucking spears, 466, 467, 470, 471
Suctorial mandibles, 453, 456
Super-Orders, 177
Supplementary Ichneumon-flies, 558
Supra-oesophageal ganglion, 117
Sutures, 92
Swarms: of locusts, 299-299 ; ; of may-flies,
441; of Termites, 362
Sympathetic nervous system, 120 ;
353
sound - instruments,
absent,
Symphrasis varia, 465
Symphyla, 42, 46, 77, 79, 80 ; structure,
61 ;
Symphyta, 503
Sympycna fusca, 415
Synaptera, 175
Synergus, 531
Syngnatha, 44
TANANA, 319
Tarachodes lucubrans, 249
Tarsus, 88, 104, 106
Taschenberg on. parthenogenesis, 141
Tausendfiisse, 41
Teeth, 95
Tegmina, 108; leaf-like, of Pterochrow,
322 ; of crickets, 331; of earwigs, 205,
212; of Phyllium, 269 : ; of Schizodacs
tylus, 325
Tegula, 103, 108
Teleganodes, 442
Telson, 205
Temples, 94
Templeton on Lepisma, 195
Tendons, 116
Tenthredinidae, 570-518
Tenthredo sp., 489 ; testes, 140
Tentorium, 99
Tepper on fossorial Blattid, 241
Terebrantia, 520
Tergum, 91, 100
Termes, 378, 380 ; 7. lucifugus, 359, 360,
364, 365, 373, 374 ; ; T. mossambicus, 356 ;
T. bellicosus, 366, 371 ; trophi, 357 ; cell
of, 367; 7. occidentis, 371; T. armiger,
371; T. tenuis, 389; T. cingulatus, 3713
T. dirus, 371; T.debilis, 371 ; T. viarum,
383
Termitarium, 386, 387 ‘
Termites, 357 f.; distinctions from ants,
502; wings, 359 ; anatomy, 360
Termitidae, 356; number of species,
389
Tertiary, 196, 216, 239, 276, 309, 340,
398, 427, 442, 449, 453, 472, 485, 533,
551, 558
Testes, 18, 49, 140, 404, 440 ; of Psocidae, —
392; of Stilopyga orientalis, 228
Tetrophthalmus chilensis, 346
Tettigides, 299, 300, 309
Tettix bipunctatus, 300
Thalessa larva, 507 ; oviposition, 554
Thamastes, 485
Thamnotrizon apterus, 316
Thecla egg, 145
Thelyotoky, 141, 498
Thermobia furnorum, 186
Thliboscelus cumellifolius, 319 i.
Thoracantha latreillei, 550 S|
Thorax, 99-103, 101, 103
Thorax porcellana wing, 227
Thyrsophorus, 395
Thysanoptera, 173
=e --”.—
‘= Pe’. - eww
INDEX 583
ee E
Thysanura, 782 f.; distinctions from Sym-
phyla, 61, 77, 79
Tibia, 88, 104’
Tillus elongatus larva, 90
Tinodes, 483
Titanophasma fayoli, 276, 428
Tomateres citrinus, 454, 458
Tomognathus, 498
- Tongue, 96—see also Lingua
Torymides, 547
Toxodera, 253; T. denticulata, 254
Trabeculae, 345
Tracheae, 128 ; absent, 553, 555
Tracheal gills, 400 f., 401—sée also
Branchiae
Tremex columba, 507
Trias, 449
Triassic, 239
Trichijulus, 76, 80
Trichodectes, 350 ; T. latus, 349
Trichoptera, 342, 473 f.
Trichostegia, 480
Tricorythus, 434, 436
Tridactylides, 340
Tridactylus variegatus, 337
Trigonalidae, 564
Trigonalys maculifrons, 564
Trigonidiides, 340
Trimen on Trachypetra bufo, 304
Trinidad, 501
Trinoton luridum, 345, 347
Trito-cerebron, 118
Trochanter, 88, 104, 491, 494, 520
Trochantin, 104; of cockroach, 222
- Trophi, 91, 94
Tryphonides, 657
Tryxalides, 303, 309, 325
Tryxalis nasuta, 279
Tubulifera, 520
Tympanophorides, 328
Tympanum, 285 f.
Tyndall on grasshopper music, 286
Uxtoa, 33
Uroceridae, 507
Useless wings, 199, 394, 484, 561
Uterus, 18, 392
‘VAGUS nervous system, 120
Van Rees on metamorphosis, 162, 164
Variation, 536 ; of colour, 252, 288, 304,
308; in desert Insects, 305; in ocelli,
267, 395, 536
Vatides, 259
Vas deferens, 18, 140, 187, 392
Vayssicre, on nymphs of Ephemeridae, 434 ;
on lingua, 438
Veins, 206
Ventral chain, 116, 187. 414; of Perlidae,
404
Ventral plate, 148 ; tube, 191, 192
Verhoeff, 38
Verlooren on circulation, 436
Vertex, 94
Vesicula seminalis, 140, 392 ; absent, 404,
414
Vespa crabro prosternum, 491
Vestibule, 112
Viallanes, on head, 87; on brain, 118,
119 ; on metamorphosis, 162
Visceral nervous system, 120
Vitellophags, 147, 152, 168
Viviparous Insects, 217, 229, 143,
218
Voetgangers, 295 f.
Vom Rath on sense organs, 122
Voracity, 250, 258
Vosseler on stink-glands, 210
WALKER, J. J., on Australian Termites,
386
Walking-leaves, 267
Walking on perpendicular and smooth
surfaces, 106
Walsh on galls, 531
Wasmann on St. Augustine’s works, 565
Wattenwyl, Brunner von—see Brunner
Weismann, on caste, 500; on meta-
morphosis, 162, 166 ; on imaginal discs,
167
Westwood, on Forficula, 204; on Helico-
mitus larva, 460, 461; on Lachesilla,
395 ; on Scleroderma, 536
Weta-punga, 326
Wheeler, on Malpighian tubes, 127 ; on
embryology of. Orthoptera, 199; on
embryology of Xiphidium, 321; on
vitellophags, 147, 152, 168; on seg-
mentation, 150
White ants, 356—-see Termites
Wielowiejski on blood-tissue, 133, 137
Will on brain of Aphididae, 118
Wingless : caddis-fly, 481; earwigs, 205 ;
Insects, 345, 352, 356, 451, 488, 536,
547 ; wingless Psocidae, 394 f.—see also
Apterous
Wings, 107 ; origin and function, 394 ; of
Blattidae, 225 f., 227; development of,
in locust, 288; in Trichoptera, 479, 480 ;
of dragon-fly, 413; of earwigs, 206 ;
of Ephemera, 431; growth of, 418;
of Ichneumon and Bracon, 559; pos-
terior absent, 466, 485; wing-hooks,
494 ; veins, 107—-see also Tegmina and
Alar Organs
** Wire-worm,” 29, 36
Wistinghausen on tracheae, 129
Wood-Mason on Cotylosoma, 272; on
mandibles, 95 ; on Mantidae, 251, 253;
on Oligotoma, 352; on phonation of
Mantidae, 258
Woodworth on embryology, 146, 153
Workers, 361, 374, 488, 495
Wyandotte Caves. Myriapods in, 34
Xptidhun eerie 321
Aye 404 494 ; metamorphosis, 110 S “pro
Xyphidviides, 507 f,510 oat Dygonterites,
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