In the case of _Hydrophilus piceus_, the {218}boat-like structure is
provided with a little mast, which is supposed by some to be for the
purpose of securing air for the eggs. _Helochares_ and _Spercheus_ (Fig.
100) carry the cocoon of eggs attached to their own bodies. _Philydrus_
constructs, one after the other, a number of these egg-bags, each
containing about fifteen eggs, and fixes each bag to the leaf of some
aquatic plant; the larvae as a rule hatch speedily, so that the advantage
of the bag is somewhat problematic.
[Illustration: Fig. 100.—_Spercheus emarginatus_ ♀. Britain. A, Upper
surface of beetle; B, under surface of abdomen, with the egg-sac ruptured
and some of the eggs escaping.]
The larvae of the aquatic division of the family have been to a certain
extent studied by Schiödte and others; those of the Sphaeridiides—the
terrestrial group of the family—are but little known. All the larvae seem
to be predaceous and carnivorous, even when the imago is of
vegetable-feeding habits; and Duméril states that in _Hydrous caraboides_
the alimentary canal undergoes a great change at the period of
metamorphosis, becoming very elongate in the adult, though in the larva it
was short. The legs are never so well developed as they are in the
Adephaga, the tarsi being merely claw-like or altogether wanting; the
mandibles are never suctorial. The respiratory arrangements show much
diversity. In most of the Hydrophilides the process is carried on by a pair
of terminal spiracles on the eighth abdominal segment, as in Dytiscidae,
and these are either exposed or placed in a respiratory chamber. In
_Berosus_ the terminal stigmata are obsolete, and the sides of the body
bear long branchial filaments. Cussac says that in _Spercheus_ (Fig. 101)
there are seven pairs of abdominal spiracles, and that the larva breathes
by presenting these to the air;[98] but Schiödte states that in this form
there are neither thoracic nor abdominal spiracles, except a pair placed in
a respiratory chamber on the eighth segment of the abdomen, after the
manner described by Miall as existing in _Hydrobius_. No doubt Cussac was
wrong in supposing the peculiar lateral abdominal processes to be
{219}stigmatiferous. In _Berosus_ there are patches of aëriferous, minute
pubescence on the body. The pupae of Hydrophilides repose on the dorsal
surface, which is protected by spinous processes on the pronotum, and on
the sides of the abdomen.
We have already remarked that this is one of the most neglected of the
families of Coleoptera, and its classification is not satisfactory. It is
usually divided into Hydrophilides and Sphaeridiides. The Sphaeridiides are
in large part terrestrial, but their separation from the purely aquatic
Hydrophilides cannot be maintained on any grounds yet pointed out.
Altogether about 1000 species of Hydrophilidae are known, but this probably
is not a tenth part of those existing. In Britain we have nearly ninety
species. Some taxonomists treat the family as a series with the name
Palpicornia. The series Philhydrida of older authors included these Insects
and the Parnidae and Heteroceridae.
[Illustration: Fig. 101—Larva of _Spercheus emarginatus_. (After
Schiödte).]
FAM. 13. PLATYPSYLLIDAE.—This consists of a single species. It will be
readily recognised from Fig. 102, attention being given to the peculiar
antennae, and to the fact that the mentum is trilobed behind. This curious
species has been found only on the beaver. It was first found by Ritsema on
American beavers (_Castor canadensis_) in the Zoological Gardens at
Amsterdam, but it has since been found on wild beavers in the Rhone in
France; in America it appears to be commonly distributed on these animals
from Alaska to Texas. It is very remarkable that a wingless parasite of
this kind should be found in both hemispheres. The Insect was considered by
Westwood to be a separate Order called Achreioptera, but there can be no
doubt that it is a beetle. It is also admitted that it shows some points of
resemblance with Mallophaga, the habits of which are similar. Its
Coleopterous nature is confirmed by the larva, which has been described by
both Horn and Riley.[99] Little is known as to the food and life-history.
Horn states that the eggs are placed on {220}the skin of the beaver amongst
the densest hair; the larvae move with a sinuous motion, like those of
Staphylinidae. It has been suggested that the Insect feeds on an Acarid,
_Schizocarpus mingaudi_; others have supposed that it eats scales of
epithelium or hairs of the beaver.
[Illustration: Fig. 102—_Platypsyllus castoris._ A, Upper side; B, lower
side, with legs of one side removed; C, antenna. (After Westwood.)]
[Illustration: Fig. 103—_Leptinus testaceus._ Britain.]
FAM. 14. LEPTINIDAE.—_Antennae rather long, eleven-jointed, without club,
but a little thicker at the extremity. Eyes absent or imperfect. Tarsi
five-jointed. Elytra quite covering abdomen. Mentum with the posterior
angles spinously prolonged._ A family of only two genera and two species.
Their natural history is obscure, but is apparently of an anomalous nature;
the inference that may be drawn from the little that is known being that
they are parasitic on mammals. There is little or nothing in their
structure to indicate this, except the condition of blindness; and until
recently the Insects were classified amongst Silphidae. _Leptinus
testaceus_ (Fig. 103) is a British Insect, and besides occurring in Europe
is well known in North America. In Europe it has been found {221}in curious
places, including the nests of mice and bumble-bees. In America it has been
found on the mice themselves by Dr. Ryder, and by Riley in the nests of a
common field-mouse, together with its larva, which, however, has not been
described. The allied genus _Leptinillus_ is said by Riley to live on the
beaver, in company with _Platypsyllus_.[100] It has been suggested that the
natural home of the _Leptinus_ is the bee's nest, and that perhaps the
beetle merely makes use of the mouse as a means of getting from one nest of
a bumble-bee to another.
FAM. 15. SILPHIDAE.—_The mentum is usually a transverse plate, having in
front a membranous hypoglottis, which bears the exposed labial palpi, and
immediately behind them the so-called bilobed ligula. The anterior coxae
are conical and contiguous: prothoracic epimera and episterna not distinct.
Visible abdominal segments usually five, but sometimes only four, or as
many as seven. Tarsi frequently five-jointed, but often with one joint
less. Elytra usually covering the body and free at the tips, but
occasionally shorter than the body, and even truncate behind so as to
expose from one to four of the dorsal plates; but there are at least three
dorsal plates in a membranous condition at the base of the abdomen._ These
beetles are extremely diverse in size and form, some being very minute,
others upwards of an inch long, and there is also considerable range of
structure. In this family are included the burying-beetles (_Necrophorus_),
so well known from their habit of making excavations under the corpses of
small Vertebrates, so as to bury them. Besides these and _Silpha_, the
roving carrion-beetles, the family includes many other very different
forms, amongst them being the larger part of the cave-beetles of Europe and
North America. These belong mostly to the genera _Bathyscia_ in Europe, and
_Adelops_ in North America; but of late years quite a crowd of these
eyeless cave-beetles of the group Leptoderini have been discovered, so that
the European catalogue now includes about 20 genera and 150 species. The
species of the genus _Catopomorphus_ are found in the nests of ants of the
genus _Aphaenogaster_ in the Mediterranean region. Scarcely anything is
known as to the lives of either the cave-Silphidae or the myrmecophilous
forms.
The larvae of several of the larger forms of Silphidae are well known, but
very little has been ascertained as to the smaller forms. {222}Those of the
burying-beetles have spiny plates on the back of the body, and do not
resemble the other known forms of the family. The rule is that the three
thoracic segments are well developed, and that ten abdominal segments are
also distinct; the ninth abdominal segment bears a pair of cerci, which are
sometimes elongate. Often the dorsal plates are harder and better developed
than is usual in Coleopterous larvae. This is especially the case with some
that are endowed with great powers of locomotion, such as _S. obscura_
(Fig. 104). The food of the larvae is as a rule decomposing animal or
vegetable matter, but some are predaceous, and attack living objects. The
larger _Silpha_ larvae live, like the _Necrophorus_, on decomposing animal
matter, but run about to seek it; hence many specimens of some of these
large larvae may sometimes be found amongst the bones of a very small dead
bird. We have found the larva and imago of _S. thoracica_ in birds' nests
containing dead nestlings. _S. atrata_ and _S. laevigata_ make war on
snails. _S. lapponica_ enters the houses in Lapland and ravages the stores
of animal provisions. _S. opaca_ departs in a very decided manner from the
habits of its congeners, as it attacks beetroot and other similar crops in
the growing state; it is sometimes the cause of serious loss to the growers
of beet. The larvae of the group Anisotomides are believed to be chiefly
subterranean in habits; that of _A. cinnamomea_ feeds on the truffle, and
the beetle is known as the truffle-beetle.
[Illustration: Fig. 104—A, Larva of _Silpha obscura_. Europe. (After
Schiödte). B, _Ptomaphila lacrymosa_, Australia.]
The number of species of Silphidae known must be at present nearer 900 than
800. Of these an unusually large proportion belong to the European and
North American regions; Silphidae being apparently far from numerous in the
tropics. Rather more than 100 species are natives of Britain. The family
reappears in considerable force in New Zealand, and is probably well
represented {223}in South Australia and Tasmania. The most remarkable form
known is perhaps the Australian genus _Ptomaphila_ (Fig. 104, B). The
classification of the family is due to Dr. Horn.[101] The only change of
importance that has since been suggested is the removal of _Sphaerites_
from this family to Synteliidae. Anisotomidae and Clambidae have been
considered distinct families, but are now included in Silphidae.
FAM. 16. SCYDMAENIDAE.—_Minute Insects allied to Silphidae, but with the
hind coxae separated, and the facets of the eyes coarser; the tarsi are
five-jointed; the number of visible abdominal segments is six._ These small
beetles are widely spread over the earth's surface, and about 700 species
are now known, of which we have about a score in Britain; many live in
ants' nests, but probably usually rather as intruders than as guests that
have friendly relations with their hosts. Nothing is known as to their
life-histories, but the food of the imago, so far as is known, consists of
Acari. _Mastigus_ is a very aberrant form, found in moss and dead leaves in
Southern Europe. By means of _Brathinus_ the family is brought very near to
Silphidae; Casey, however, considers _Brathinus_ to belong to Staphylinidae
rather than to Scydmaenidae. The South European _Leptomastax_ is remarkable
on account of the slender, long, sickle-shaped mandibles. The Oriental
genus _Clidicus_ is the largest and most remarkable form of the family; it
has a very slender neck to its broad head, and is more than a quarter of an
inch long.
FAM. 17. GNOSTIDAE.—_Minute Insects with three-jointed antennae,
five-jointed tarsi, and three apparent ventral segments, the first of
which, however, is elongate, and consists of three united plates. Elytra
entirely covering the after-body._ The family consists of two species which
have been found in the nests of ants, of the genus _Cremastogaster_, in
Brazil.[102]
FAM. 18. PSELAPHIDAE.—_Very small Insects; the elytra much abbreviated,
usually leaving as much as half the abdomen uncovered; the maxillary palpi
usually greatly developed, and of a variety of remarkable forms; the
segments of the abdomen not more than seven in number, with little or no
power of movement. Tarsi with not more than three joints._ These small
Coleoptera mostly live in the nests of ants, and present a great diversity
of extraordinary {224}shapes, and very peculiar structures of the antennae
and maxillary palpi. Owing to the consolidation of some of its segments,
the abdomen frequently appears to have less than the usual number. In the
curious sub-family Clavigerides, the antennae may have the joints reduced
to two or even, to all appearance, to one; the tarsi suffer a similar
reduction. There are about 2500 species of Pselaphidae known; many of them
have never been found outside the ants' nests; very little, however, is
known as to their natural history. It is certain that some of them excrete,
from little tufts of peculiar pubescence, a substance that the ants are
fond of. The secretory patches are found on very different parts of the
body and appendages. _Claviger testaceus_ is fed by the ants in the same
way as these social Insects feed one another; the _Claviger_ has also been
seen to eat the larvae of the ants. They ride about on the backs of the
ants when so inclined. The family is allied to Staphylinidae, but is easily
distinguished by the rigid abdomen. Only one larva—that of _Chennium
bituberculatum_—is known. It appears to be very similar to the larvae of
Staphylinidae. The best account of classification and structure is that
given by M. Achille Raffray,[103] who has himself discovered and described
a large part of the known species.
FAM. 19. STAPHYLINIDAE.—_Elytra very short, leaving always some of the
abdominal segments exposed, and covering usually only two of the segments.
Abdomen usually elongate, with ten dorsal, and seven or eight ventral
segments; of the latter six or seven are usually exposed; the dorsal plates
as hard as the ventral, except sometimes in the case of the first two
segments; the segments very mobile, so that the abdomen can be curled
upwards. The number of tarsal joints very variable, often five, but
frequently as few as three, and not always the same on all the feet._
Staphylinidae (formerly called Brachelytra or Microptera) is one of the
most extensive of even the great families of Coleoptera; notwithstanding
their diversity, they may in nearly all cases be recognised by the more
than usually mobile and uncovered abdomen, combined with the fact that the
parts of the mouth are of the kind we have mentioned in Silphidae. The
present state of the classification of this family has been recently
discussed by Ganglbauer.[104]
{225}[Illustration: Fig. 105—Staphylinidae. A, Larva of _Philonthus
nitidus_. Britain. (After Schiödte.) B, _Ocypus olens_, Britain; C, tip of
abdomen, of _O. olens_ with stink-vessels.]
At present about 9000 species are known, some of which are minute, while
scarcely any attain a size of more than an inch in length, our common
British black cock-tail, or "devil's coach-horse beetle," _Ocypus olens_,
being amongst the largest. Though the elytra are short, the wings in many
forms are as large as those of the majority of beetles; indeed many
Staphylinidae are more apt at taking flight than is usual with Coleoptera;
the wings when not in use are packed away under the short elytra, being
transversely folded, and otherwise crumpled, in a complicated but orderly
manner. It is thought that the power of curling up the abdomen is connected
with the packing away of the wings after flight; but this is not the case:
for though the Insect sometimes experiences a difficulty in folding the
wings under the elytra after they have been expanded, yet it overcomes this
difficulty by slight movements of the base of the abdomen, rather than by
touching the wings with the tip. What the value of this exceptional
condition of short elytra and corneous dorsal abdominal segments to the
Insect may be is at present quite mysterious. The habits of the members of
the family are very varied; many run with great activity; the food is very
often small Insects, living or dead; a great many are found in fungi of
various kinds, and perhaps eat them. It is in this family that we meet with
some of the most remarkable cases of symbiosis, i.e. lives of two kinds of
creatures mutually accommodated with good will. The relations between the
Staphylinidae of the genera _Atemeles_ and _Lomechusa_, and certain ants,
in the habitations of which they dwell, are very interesting. The beetles
are never found out of the ants' nests, or at any rate not very far from
them. The most friendly relations exist between them and the ants: they
have patches of yellow hairs, and these {226}apparently secrete some
substance with a flavour agreeable to the ants, which lick the beetles from
time to time. On the other hand, the ants feed the beetles; this they do by
regurgitating food, at the request of the beetle, on to their lower lip,
from which it is then taken by the beetle (Fig. 82). The beetles in many of
their movements exactly resemble the ants, and their mode of requesting
food, by stroking the ants in certain ways, is quite ant-like. So
reciprocal is the friendship that if an ant is in want of food, the
_Lomechusa_ will in its turn disgorge for the benefit of its host. The
young of the beetles are reared in the nests by the ants, who attend to
them as carefully as they do to their own young. The beetles have a great
fondness for the ants, and prefer to sit amongst a crowd thereof; they are
fond of the ants' larvae as food, and indeed eat them to a very large
extent, even when their own young are receiving food from the ants. The
larva of _Lomechusa_, as described by Wasmann (to whom we are indebted for
most of our knowledge of this subject),[105] when not fully grown, is very
similar to the larvae of the ants; although it possesses legs it scarcely
uses them: its development takes place with extraordinary rapidity, two
days, at most, being occupied in the egg, and the larva completing its
growth in fourteen days. Wasmann seems to be of opinion that the ants
scarcely distinguish between the beetle-larvae and their own young; one
unfortunate result for the beetle follows from this, viz. that in the pupal
state the treatment that is suitable for the ant-larvae does not agree with
the beetle-larvae: the ants are in the habit of digging up their own kind
and lifting them out and cleaning them during their metamorphosis; they
also do this with the beetle-larvae, with fatal results; so that only those
that have the good fortune to be forgotten by the ants complete their
development. Thus from thirty _Lomechusa_ larvae Wasmann obtained a single
imago, and from fifty _Atemeles_ larvae not even one.
Many other Staphylinidae are exclusively attached to ants' nests, but most
of them are either robbers, at warfare with the ants—as is the case with
many species of _Myrmedonia_ that lurk about the outskirts of the nests—or
are merely tolerated by the ants, not receiving any direct support from
them. The most {227}remarkable Staphylinidae yet discovered are some
viviparous species, forming the genera _Corotoca_ and _Spirachtha_, that
have very swollen abdomens, and live in the nests of Termites in
Brazil:[106] very little is, however, known about them. A very large and
powerful Staphylinid, _Velleius dilatatus_, lives only in the nests of
hornets and wasps. It has been supposed to be a defender of the
Hymenoptera, but the recent observations of Janet and Wasmann make it clear
that this is not the case: the _Velleius_ has the power of making itself
disagreeable to the hornets by some odour, and they do not seriously attack
it. The _Velleius_ finds its nutriment in larvae or pupae of the wasps that
have fallen from their cells, or in other organic refuse.
The larvae of Staphylinidae are very similar to those of Carabidae, but
their legs are less perfect, and are terminated only by a single claw;
there is no distinct labrum. The pupae of some are obtected, _i.e._ covered
by a secondary exudation that glues all the appendages together, and forms
a hard coat, as in Lepidoptera. We have about 800 species of Staphylinidae
in Britain, and it is probable that the family will prove one of the most
extensive of the Order. It is probable that one hundred thousand species or
even more are at present in existence.
FAM. 20. SPHAERIIDAE.—_Very minute. Antennae eleven-jointed, clubbed. Tarsi
three-jointed. Abdomen with only three visible ventral segments_. This
family includes only three or four species of Insects about 1/50 of an inch
long. They are very convex, and be found walking on mud. _S. acaroides_
occurs in our fens. Mr. Matthews considers that they are most nearly allied
to Hydrophilidae.[107]
[Illustration: Fig. 106—_Trichopteryx fascicularis_. Britain. A, Outline of
perfect Insect; B, part of upper surface; C, larva from side; D, from
above; E, pupa; F, wing; G, natural size of imago.]
FAM. 21. TRICHOPTERYGIDAE.—_Extremely minute: antennae {228}clavicorn
(basal and apical joints thicker than middle joints); tarsi three-jointed;
elytra sometimes covering abdomen, in other cases leaving a variable number
of segments exposed; wings fringed._ This family comprises the smallest
Insects; _Nanosella fungi_ being only 1/100 of an inch long, while the
largest Trichopterygid is only 1/12 of an inch. The small size is not
accompanied by any degeneration of structure, the minute, almost invisible
forms, having as much anatomical complexity as the largest Insects. Very
little is known as to the natural history. Probably these Insects exist in
all parts of the world, for we have about eighty species in England, and
Trichopterygidae are apparently numerous in the tropics.[108]
FAM. 22. HYDROSCAPHIDAE.—_Extremely minute aquatic Insects, with elongate
abdomen. Antennae eight-jointed._ The other characters are much the same as
those we have mentioned for Trichopterygidae. The family is not likely to
come before the student, as only three or four species from Southern Europe
and North America are known.[109]
[Illustration: Fig. 107—A, Larva of _Orthoperus brunnipes_ (after Perris);
B, _O. atomarius_, perfect Insect. Britain.]
FAM. 23. CORYLOPHIDAE.—_Minute beetles. Tarsi four-jointed, but appearing
only three-jointed, owing to the hind joint being concealed by the
emarginate (or notched) second joint. Six free ventral segments. Maxillae
with only one lobe. Antennae of peculiar form._ There are about 200 species
of these little Insects, but the family is apparently represented all over
the world, and will probably prove to be much more extensive. The peculiar
larva of _Orthoperus brunnipes_ was found abundantly by Perris in thatch in
France. Mr. Matthews proposes to separate the genus _Aphanocephalus_ as a
distinct family, Pseudocorylophidae.[110] In Corylophidae the wings are
fringed with long hairs, as is the case in so many small Insects: the
species of _Aphanocephalus_ are rather larger Insects, and the wings are
not fringed; the tarsi are only three-jointed.
{229}FAM. 24. SCAPHIDIIDAE.—_Front coxae small, conical; prothorax very
closely applied to the after-body; hind coxae transverse, widely separated:
abdomen with six or seven visible ventral plates; antennae at the extremity
with about five joints that become gradually broader. Tarsi five-jointed._
This family consists of a few beetles that live in fungi, and run with
extreme rapidity; they are all small, and usually rare in collections. Some
of the exotic forms are remarkable for the extreme tenuity and fragility of
the long antennae, which bear fine hairs. The number of described species
does not at present reach 200, but the family is very widely distributed.
We have three or four species in Britain. All we know of the larvae is a
description of that of _Scaphisoma agaricinum_ by Perris;[111] it is like
the larva of Staphylinidae, there are nine abdominal segments in addition
to a very short, broad pseudopod, and very short cerci. This larva feeds on
agarics; it goes through its development in about three weeks; unlike the
adult it is not very active.
[Illustration: Fig. 108—_Scaphisoma agaricinum._ Britain. A Larva (after
Perris); B perfect Insect.]
FAM. 25. SYNTELIIDAE.—_Antennae clavicorn, with very large club: labium,
with hypoglottis and the parts beyond it, exposed. Front coxae transverse.
Abdomen with five visible ventral segments, and eight or nine dorsal, the
basal four of which are semi-corneous._ This family includes only five
species; its classification has given rise to much difference of opinion.
We have, after consideration of all its characters, established it as a
distinct family[112] allied to Silphidae. The perfect Insects live on the
sap running from trees: but nothing else is known of their natural history.
Like so many others of the very small families of aberrant Coleoptera, it
has a very wide distribution; _Syntelia_ being found in Eastern Asia and
Mexico, while the sub-family Sphaeritides occurs, as a single species, in
Europe and North America. The earlier instars are unknown.
[Illustration: Fig. 109—_Syntelia westwoodi._ Mexico. (From _Biol. Centr.
Amer._)]
{230}[Illustration: Fig. 110—_Platysoma depressum._ Europe. A, Larva (after
Schiödte); B, perfect Insect.]
FAM. 26. HISTERIDAE.—_Very compact beetles, with very hard integument,
short, bent antennae, with a very compact club: no hypoglottis. Elytra
closely applied to body, but straight behind, leaving two segments exposed.
Abdomen with five visible ventral segments; with seven dorsal segments, all
hard. Front coxae strongly transverse, hind coxae widely separated._ The
extremely compact form, and hard integument, combined with the peculiar
antennae—consisting of a long basal joint, six or seven small joints, and
then a very solid club of three joints covered with minute
pubescence—render these Insects unmistakable. The colour is usually shining
black, but there are numerous departures from this. The way in which these
Insects are put together so as to leave no chink in their hard exterior
armour when in repose, is very remarkable. The mouth-parts are rather
highly developed, and the family is entitled to a high rank; it consists at
present of about 2000 species;[113] in Britain we have about 40. The larvae
are without ocelli or labrum, but have well-developed mandibles, the second
and third thoracic segments being short, the ninth segment of the abdomen
terminal, with two distinctly jointed cerci.[114] Histeridae are common in
dung, in carcases, decaying fungi, etc., and some live under bark—these
being, in the case of the genus _Hololepta_, very flat. Some are small
cylinders, elaborately constructed, for entering the burrows of Insects in
wood (_Trypanaeus_); a certain number are peculiar to ants' nests. Formerly
it was supposed that the Insects were nourished on the decaying substances,
but it is now believed, with good reason, that they are eminently
predaceous, in both larval and imaginal {231}instars, and devour the larvae
of Diptera, etc. The relations of the ants'-nest forms to the ants is not
made out, but it is highly probable that they eat the ants' larvae, and
furnish the ants with some dainty relish. A few species live in company
with Termites.
FAM. 27. PHALACRIDAE.—_Body very compact; elytra entirely covering it;
apical joints of antennae rather broader, usually long; front coxae
globular; posterior coxae contiguous; abdomen with five visible ventral
segments; tarsi five-jointed, fourth joint usually small and obscure._ This
family consists entirely of small Insects: the tarsal structure is very
aberrant, and is also diverse, so that the student may without careful
observation pass the Insects over as having only four-jointed tarsi; their
structure, so far as the front two pairs are concerned, being very nearly
that of many Phytophaga. The larvae live in the heads of flowers,
especially of the flowers of Compositae. Having bored their way down the
stems, they pupate in earthen cocoons. Heeger[115] says that he has
observed in favourable seasons six generations; but the larvae die readily
in unfavourable seasons, and are destroyed in vast numbers when the meadows
are mowed. Seven years ago very little was known as to the family, and the
list of their species scarcely amounted to 100, but now probably 300 are
described. They occur in all parts of the world; we have fourteen in
Britain.
[Illustration: Fig. 111—_Olibrus bicolor._ Europe. A, Larva (after Heeger);
B, perfect Insect.]
FAM. 28. NITIDULIDAE.—_Antennae with a three-jointed club; all the coxae
separated, and each with an external prolongation; tarsi five-jointed, the
fourth joint smaller than any of the others; abdomen with five visible
plates._ These Insects are numerous, about 1600 species being at present
known; in many of them the elytra nearly or quite cover the hind body, but
in many others they are more or less abbreviated; in this case the Insects
may be distinguished from Staphylinidae by the form of their antennae, and
the smaller number of visible ventral segments. The habits are very varied,
a great many are found on flowers, others are attracted by the sap of
trees; some live in carcases. We have about 90 species in Britain; several
forms of {232}the genera _Meligethes_ and _Epuraea_ are among the most
abundant of our beetles. Most of what is known as to the larvae is due to
Perris; several have been found living in flowers; that of _Pria_ haunts
the flower of _Solanum dulcamara_ at the junction of the stamens with the
corolla; the larva of _Meligethes aeneus_ sometimes occasions much loss by
preventing the formation of seed in cultivated Cruciferae, such as Rape.
These floricolous larvae grow with great rapidity, and then leave the
flowers to pupate in the ground. The larva of _Nitidula_ lives in carcases,
though it is not very different from that of _Pria_. The larva of _Soronia_
lives in fermenting sap, and has four hooks curving upwards at the
extremity of the body. The curious genus _Cybocephalus_ consists of some
very small, extremely convex Insects that live in flowers in Southern
Europe; they have only four joints to the tarsi. The perfect Insects of the
group Ipides are remarkable from having a stridulating organ on the front
of the head. The classification of the well-known genus _Rhizophagus_ has
given rise to much discussion; although now usually placed in Nitidulidae,
we think it undoubtedly belongs to Cucujidae.
[Illustration: Fig. 112—_Pria dulcamarae._ Britain. A, Larva (after
Perris); B, perfect Insect.]
[Illustration: Fig. 113—_Temnochila coerulea._ Europe. A, Larva (after
Perris); B, perfect Insect.]
FAM. 29. TROGOSITIDAE.—_Differs from Nitidulidae in the structure of the
tarsi; these appear to be four-jointed, with the third joint similar in
size and form to the preceding; they are, however, really five-jointed, an
extremely short basal joint being present. Hind coxae contiguous. The club
of each antenna is bilaterally asymmetric, and the sensitive surface is
confined to certain parts of the joints._ There are some 400 or 500 species
of Trogositidae, but nearly all of them are exotic. The larvae (Fig. 113,
A), are predaceous, destroying other larvae in large numbers, and it is
probable that the imagos do the same. The larva of _Tenebroides_ (better
known as _Trogosita_) _mauritanica_ is found in corn and flour, and is said
to have sometimes been very {233}injurious by eating the embryo of the
corn, but it is ascertained that it also devours certain other larvae that
live on the corn. This beetle has been carried about by commerce, and is
now nearly cosmopolitan. Our three British species of Trogositidae
represent the three chief divisions of the family, viz. Nemosomides,
Temnochilides, Peltides; they are very dissimilar in form, the Peltides
being oval, with retracted head. It is doubtful whether the members of the
latter group are carnivorous in any of their stages; it is more probable
that they live on the fungi they frequent. Peltidae stand as a distinct
family in many works.[116]
[Illustration: Fig. 114—_Bitoma crenata._ Britain. A, Larva (after Perris);
B, perfect Insect.]
FAM. 30. COLYDIIDAE.—_Antennae with a terminal club, tarsi four-jointed,
none of the joints broad; front and middle coxae small, globose, embedded;
hind coxae transverse, either contiguous or separated; five visible ventral
segments, several of which have no movement._ This is a family of interest,
owing to the great diversity of form, to the extraordinary sculpture and
clothing exhibited by many of its members, and to the fact that most of its
members are attached to the primitive forests, and disappear entirely when
these are destroyed. We have fifteen species in Britain, but about half of
them are of the greatest rarity. There are about 600 species known at
present; New Zealand has produced the greatest variety of forms; the
forests of Teneriffe are rich in the genus _Tarphius_. The sedentary lives
of many of these beetles are very remarkable; the creatures concealing
themselves in the crannies of fungus-covered wood, and scarcely ever
leaving their retreats, so that it is the rarest circumstance to find them
at any distance from their homes. _Langelandia anophthalma_ lives entirely
underground and is quite blind, the optic lobes being absent. Some
Colydiidae are more active, and enter the burrows of wood-boring Insects to
destroy the larvae (_Colydium_). Few of the larvae are known; but all
appear to have the body terminated by peculiar hard corneous processes, as
is the case with a great variety of Coleopterous larvae that live in
wood.[117]
{234}FAM. 31. RHYSODIDAE.—_Tarsi four-jointed; mouth-parts covered by the
large mentum; front tibiae notched on the inner edge._ This family consists
only of a few species, but is found nearly all over the world in the warm
and temperate regions. In many of their characters they resemble the
Adephaga, but are very different in appearance and in the mouth. The larvae
are not known. Some authorities think these Insects should be placed in the
series Adephaga,[118] but it is more probable that they will prove to be
amongst the numerous aberrant forms of Coleoptera that approach the various
large natural series, without really belonging to them. The three families,
Colydiidae, Cucujidae, and Rhysodidae, exhibit relations not only with
other families of the Coleoptera Polymorpha, but also with most of the
great series; Adephaga, Rhynchophora, Phytophaga, and Heteromera, being
each closely approached.
FAM. 32. CUCUJIDAE.—_Tarsi five- or four-jointed, the first joint often
short: antennae sometimes clubbed, but more often quite thin at the tip;
front and middle coxae deeply embedded, globular, but with an angular
prolongation externally; abdomen with five visible ventral segments, all
movable._ This family and the Cryptophagidae are amongst the most difficult
families to define; indeed it is in this portion of the Clavicorns that an
extended and thorough study is most urgently required. The Cucujidae
include a great diversity of forms; they are mostly found under the bark of
trees, and many of them are very flat. Many of the larvae are also very
flat, but Ferris says there is great diversity in their structure: they are
probably chiefly carnivorous. There are about 400 species described; we
have nearly a score in Britain.
[Illustration: Fig. 115—_Brontes planatus._ Britain. A, Larva; B, pupa; C,
perfect Insect. (A and B after Perris.)]
The family Cupesidae of certain taxonomists must be at present associated
with Cucujidae, though the first joint of the tarsus is elongate.
{235}FAM. 33. CRYPTOPHAGIDAE.—_Front and middle coxae very small and deeply
embedded; antennae with enlarged terminal joints; tarsi five-jointed, the
posterior sometimes in the male only four-jointed; abdomen with five
visible ventral segments, capable of movement, the first much longer than
any of the others._ A small family composed of obscure forms of minute
size, which apparently have mould-eating habits, though very little is
known on this point and several of the genera (_Antherophagus_,
_Telmatophilus_) are found chiefly on growing plants, especially in
flowers. Although the imago of _Antherophagus_ lives in flowers, yet the
larva has only been found in the nests of bumble-bees; there is reason for
believing that the imago makes use of the bee to transport it from the
flowers it haunts to the nests in which it is to breed;[119] this it does
by catching hold of the bee with its mandibles when the bee visits the
flower in which the beetle is concealed. It is strange the beetle should
adopt such a mode of getting to its future home, for it has ample wings. We
must presume that its senses and instinct permit it to recognise the bee,
but do not suffice to enable it to find the bee's nest. Some of the larvae
of the genus _Cryptophagus_ are found abundantly in the nests of various
wasps, where they are probably useful as scavengers, others occur in the
nests of social caterpillars, and they are sometimes common in loose straw;
this being the habitat in which Perris found the one we figure.
[Illustration: Fig. 116—_Cryptophagus dentatus._ Britain. A, Larva (after
Perris); B, perfect Insect.]
FAM. 34. HELOTIDAE.—_Front and middle coxal cavities round, with scarcely
any angular prolongation externally; all the coxae widely separated; five
visible ventral segments, all mobile._ The Insects of this family are
closely allied to Trogositidae and Nitidulidae, and have the tarsal
structure of the former family; but the Helotidae are different in
appearance from any members of either of these two families, and are
readily distinguished by the coxal character. They are frequently
classified with the Erotylidae, from which they differ by the differently
shaped feet, especially by the diminished basal joint. {236}There is but
one genus, and for a long time only two or three species were known, and
were great rarities in collections; in the last few years the number has
been raised to nearly forty.[120] They are remarkable beetles with oblong
form, and a somewhat metallic upper surface, which is much sculptured, and
possesses four yellow, smooth spots on the elytra. According to Mr. George
Lewis they are found feeding at the running sap of trees, but the larvae
are not known. Helotidae are peculiar to the Indo-Malayan region (including
Japan) with one species in Eastern Africa.
FAM. 35. THORICTIDAE.—_Tarsi five-jointed, none of the joints broad; front
coxae small, rather prominent, but not at all transverse; five visible
ventral plates, all mobile; metasternum very short; antennae short, with a
solid club._ This little family, consisting of the genus _Thorictus_,
appears to be a distinct one, though the structure has only been very
imperfectly studied. It is peculiar to the Mediterranean region, where the
species live in ants' nests. They appear to be on terms of great intimacy
with the ants; a favourite position of the beetle is on the scape of the
antenna of an ant; here it hooks itself on firmly, and is carried about by
the ant. Like so many other ants'-nest beetles, Thorictidae possess tufts
of golden hair, which secrete some substance, the flavour of which is
appreciated by the ants; these tufts in Thorictidae are situated either at
the hind angles of the pronotum, or on the under surface of the body on
each side of the breast; Wasmann thinks that when the beetles are riding
about, as above described, the ants have then an opportunity of getting at
the patches on the under surface.
[Illustration: Fig. 117—_Tritoma bipustulata._ Erotylidae. Britain. A,
Larva (after Perris); B, perfect Insect.]
FAM. 36. EROTYLIDAE.—_Tarsi five-jointed, but with the fourth usually very
small, the first three more or less broad and pubescent beneath. Antennae
strongly clubbed. Front and middle coxal acetabula round, without angular
prolongation externally; five visible ventral segments._ This is now a
large and important family of about 1800 species, but it is chiefly exotic
and tropical, its members haunting the fungoid growths {237}in forests. We
have only six species in Britain, and the whole of Europe has only about
two dozen, most of them insignificant (and in the case of the Dacnides
aberrant, approaching the Cryptophagidae very closely). The sub-family
Languriides (quite wanting in Europe) consists of more elongate Insects,
with front acetabula open behind; they have different habits from
Erotylides proper; some are known to live as larvae in the stems of
herbaceous plants. They possess a highly developed stridulating organ on
the front of the head. The Clavicorn Polymorpha are very closely connected
with the Phytophaga by Languriides.
FAM. 37. MYCETOPHAGIDAE.—_Tarsi four-jointed, slender, the front feet of
the male only three-jointed; coxae oval, not deeply embedded; abdomen with
five ventral segments, all movable._ A small family, of interest chiefly
because of the anomaly in the feet of the two sexes, for which it is
impossible to assign any reason. The species are small, uninteresting
Insects that live chiefly on Cryptogams of various kinds, especially in
connection with timber; the larvae being also found there. There are about
a dozen species in Britain, and scarcely 100 are described from all the
world. The Diphyllides, placed by Leconte and Horn in this family, seem to
go better in Cryptophagidae.
[Illustration: Fig. 118—_Litargus bifasciatus._ Mycetophagidae. Britain. A,
Larva (after Perris); B, perfect Insect.]
FAM. 38. COCCINELLIDAE (_Lady-birds_).—_Tarsi apparently three-jointed; the
first two joints pubescent beneath; the third joint consisting really of
two joints, the small true third joint being inserted near the base of the
second joint, the upper surface of which is grooved to receive it. Head
much concealed by the thorax. Antennae feebly clubbed._ The lady-birds
number fully 2000 species. The structure of their feet distinguishes them
from nearly all other Coleoptera except Endomychidae, which are much less
rotund in form, and have larger antennae. One genus of
Endomychids—_Panomoea_—bears, however, a singular resemblance to
lady-birds, both in form and style of coloration. Several species of
Coccinellidae are remarkable on account of the numerous variations in
colour they present. Coccinellidae frequently multiply to an enormous
extent, and are of great value, as they destroy wholesale the plant-lice,
scale-Insects, {238}and Acari that are so injurious to cultivated plants.
They also eat various other soft-bodied Insects that attack plants. As they
are excessively voracious, and are themselves singularly free from enemies
and multiply with great rapidity, all these features of their economy
render them of inestimable value to the agriculturist and horticulturist.
The species of the sub-family Epilachnides feed on plants, and one or two
are occasionally injurious. The body-fluid of Coccinellidae has an
unpleasant odour and taste. Many lady-birds have the power of exuding, when
disturbed, small quantities of a yellow fluid. Lutz has shown that this is
not a special secretion, but an exudation of the fluid of the body that
takes place through a small orifice at the tip of the tibia, from pressure
caused by contraction of the body and limb.[121]
The larvae are much more active than beetle-larvae usually are, and many of
them are very conspicuous when running about on plants to hunt their prey.
They usually cast their skins three times, and sometimes concomitantly
change a good deal in colour and form; the larval life does not usually
exceed four or five weeks; at the end of which time the larva suspends
itself by the posterior extremity, which is glued by a secretion to some
object; the larval skin is pushed back to the anal extremity, disclosing
the pupa; this differs in several respects from the usual pupa of beetles;
it is harder, and is coloured, frequently conspicuously spotted, and
dehisces to allow the escape of the beetle, so that the metamorphosis is
altogether more like that of Lepidoptera than that of Coleoptera. There is
much variety in the larvae; some of them bear large, complexly-spined,
projections; those of the group Scymnites have small depressions on the
surface, from which it has been ascertained that waxy secretions exude; but
in _Scymnus minimus_ no such excretions are formed. Certain species, when
pupating, do not shuffle the skin to the extremity of the body, but retain
it as a covering for the pupa. The larvae that feed on plants are much less
active than the predaceous forms. We are well supplied with Coccinellidae
in Britain, forty species being known here.
The systematic position of Coccinellidae amongst the Coleoptera has been
for long a moot point. Formerly they were associated with various other
beetles having three-jointed, or apparently three-jointed, feet, as a
series with the name Trimera, or {239}Pseudotrimera. But they are generally
placed in the Clavicorn series, near Endomychidae. Verhoeff has recently
made considerable morphological studies on the male genital organs of
Coleoptera, and as the result, he concludes that Coccinellidae differ
radically from all other Coleoptera as regards these structures, and he
therefore treats them as a distinct series or sub-order, termed
Siphonophora. The genus _Lithophilus_ has been considered doubtfully a
member of Coccinellidae, as the tarsi possess only in a slight degree the
shape characteristic of the family: Verhoeff finds that they are truly
Coccinellidae, forming a distinct division, Lithophilini; and our little
species of _Coccidula_, which have somewhat the same appearance as
Lithophilini, he treats as another separate group, Coccidulini.
FAM. 39. ENDOMYCHIDAE.[122]—_Tarsi apparently three-jointed, the first two
joints broad, the terminal joint elongate; at the base of the terminal
joint there is, however, a very small joint, so that the tarsi are
pseudotetramerous; antennae rather large, with a large club; labium not at
all retracted behind the mentum; front and middle coxae globose; abdomen
with five movable ventral segments, and a sixth more or less visible at the
tip._ This family includes a considerable diversity of elegant Insects that
frequent fungoid growths on wood. It comprises at present fully 500
species, but nearly the whole of them are exotic, and inhabit the tropical
forests. We have only two British species, both of which are now rarities,
but apparently were much commoner at the beginning of the century. The
larvae are broader than is usual in Coleoptera; very few, however, are
known.
[Illustration: Fig. 119—_Mycetaea hirta._ Britain. A, Larva (after
Blisson); B, perfect Insect.]
FAM. 40. MYCETAEIDAE.—_Tarsi four-jointed, the first two joints not very
different from the third, usually slender; abdomen with five visible
ventral segments, which are movable; front and middle coxae globular._ The
little Insects composing this family are by many placed as a division of
Endomychidae, and Verhoeff is of opinion that the group is an altogether
artificial one; but we think, with Duval, it makes matters simpler to
separate them. There are only {240}some forty or fifty species, found
chiefly in Europe and North America. We have three in Britain; one of
these, _Mycetaca hirta_ is very common, and may be found in abundance in
cellars in the heart of London, as well as elsewhere; it is said to have
injured the corks of wine-bottles, and to have caused leakage of the wine,
but we think that it perhaps only increases some previous deficiency in the
corkage, for its natural food is fungoid matters. The larva is remarkable
on account of the clubbed hairs at the sides of the body.
FAM. 41. LATRIDIIDAE.—_Tarsi three-jointed; anterior coxal cavities round,
not prolonged externally; abdomen with five visible and mobile ventral
segments._ Very small Insects, species of which are numerous in most parts
of the world, the individuals being sometimes very abundant. The larvae
(Fig. 120, A) are said by Perris to have the mandibles replaced by fleshy
appendages. The pupa of _Latridius_ is remarkable, on account of the
numerous long hairs with heads instead of points; the larva of _Corticaria_
is very like that of _Latridius_, but some of the hairs are replaced by
obconical projections. The sub-family Monotomides is by many treated as a
distinct family; they have the elytra truncate behind, exposing the
pygidium, and the coxae are very small and very deeply embedded. Most of
the Latridiidae are believed to live on fungoid matters; species of
_Monotoma_ live in ants' nests, but probably have no relations with the
ants. A few species of Latridiides proper also maintain a similar life;
_Coluocera formicaria_ is said to be fond of the stores laid up by
_Aphaenogaster structor_ in its nests. About 700 species are now known;
scarcely any of the individuals are more than one-tenth of an inch long. We
have about 40 species in Britain. The North American genus _Stephostethus_
has the prosternum constructed behind the coxae, somewhat in the same
manner as it is in the Rhynchophorous series of Coleoptera.
[Illustration: Fig. 120—_Latridius minutus._ Britain. A, Larva (after
Perris); B, perfect Insect.]
FAM. 42. ADIMERIDAE.—_Tarsi appearing only two-jointed, a broad basal joint
and an elongate claw-bearing joint, but between the two there are two very
small joints._ This family consists only of the American genus _Adimerus_;
nothing is known of {241}the life-history of these small Insects. They are
of some interest, as this structure of the foot is not found in any other
beetles.
[Illustration: Fig. 121—_Adimerus setosus._ Adimeridae. A, the Insect; B,
one foot more enlarged. Mexico. From _Biol. Centr. Amer. Col._ ii. pt. i.]
[Illustration: Fig. 122—_Tiresias serra._ Larva. New Forest.]
FAM. 43. DERMESTIDAE.—_Tarsi five-jointed; antennae usually short, with the
club frequently very large in proportion, and with the under side of the
thorax bearing a hollow for its reception. Front coxae rather long,
oblique: hind coxa formed to receive the femur when in repose._ A family of
300 or 400 species of small or moderate-sized beetles; the surface, usually
covered with fine hair, forming a pattern, or with scales. _Byturus_, the
position of which has long been disputed, has now been placed in this
family; it has a more imperfectly formed prosternum, and the third and
fourth joints of the tarsi are prolonged as membranous lobes beneath; the
hind coxae leave the femora quite free. Dermestidae in the larval state
nearly all live on dried animal matter, and are sometimes very destructive;
some of them totally destroy zoological collections. They are very
remarkable on account of the complex clothing of hairs they bear; they have
good powers of locomotion, and many of them have a peculiar gait, running
for a short distance, then stopping and vibrating some of their hairs with
extreme rapidity. They exhibit great variety of form. Many of them are
capable of supporting life for long periods on little or no food, and in
such cases moult an increased number of times: pupation takes place in the
larval skin. _Anthrenus fasciatus_ has been reared in large numbers on a
diet of dried horse-hair in furniture. The young larva of this species
observed by the writer did not possess the remarkable, complex arrangement
of hairs that appeared when it was further grown. The most curious of
Dermestid larvae is that of _Tiresias serra_, which lives amongst cobwebs
in {242}old wood, and probably feeds on the remains of Insects therein,
perhaps not disdaining the cobwebs themselves. Attention has been
frequently called to the hairs of the larvae of these Insects, but they
have never been adequately discussed, and their function is quite unknown.
FAM. 44. BYRRHIDAE (_Pill-beetles_).—_Oval or round, convex beetles; tarsi
five-jointed, front coxae not exserted, transverse; hind coxa shielding the
retracted femur. The whole of the appendages capable of a complete
apposition to the body._ Although a small family of only 200 or 300
species, Byrrhidae are so heterogeneous that no characteristic definition
that will apply to all the sub-families can be framed. Very little is known
as to their life-histories. _Byrrhus pilula_ is one of our commonest
beetles, and may be found crawling on paths in early spring even in towns;
it moves very slowly, and when disturbed, at once contracts the limbs so
completely that it looks like an inanimate object. The larva is
cylindrical, soft; the prothoracic and last two abdominal segments are
larger than the others, the last bearing two pseudopods; its habits are
unknown, and no good figure exists of it.
The chief groups of Byrrhidae are Nosodendrides, Byrrhides (including
Amphicyrtides), Limnichides, and Chelonariides. The first consists of
species frequenting the exuding sap of trees; they have an unusually large
mentum, abruptly clubbed antennae, and the head cannot be retracted and
concealed. The genus _Nosodendron_ seems to be distributed over a large
part of the world. The Byrrhides have the antennae gradually thicker
towards the tip, the mentum small, and the head and thorax so formed that
the former can be perfectly retracted. The species are rather numerous, and
are found in the northern and antipodeal regions, being nearly completely
absent from the tropics. The Limnichides are minute Insects living in very
moist places; they have small delicate antennae, which are imperfectly
clubbed. The group is very widely distributed.
The Chelonariides are a very peculiar form of Coleoptera: oval Insects of
small size with the prothorax so formed that the head can be withdrawn
under (rather than into) it, and then abruptly inflexed, so that the face
then forms part of the under surface: the antennae have the basal three
joints thicker than the others; these being not in the least clubbed, but
having the {243}joints so delicately connected that the organs are rarely
unmutilated. The modifications of the head and prothorax are quite unlike
those of other Byrrhidae, and if the Chelonariides do not form a distinct
family, they should be associated with Dascillidae. Nothing is known as to
the earlier stages. They are chiefly tropical Insects, though one species
is found in North America.
FAM. 45. CYATHOCERIDAE.—_Minute Insects of broad form; parts of the mouth
concealed; antennae four-jointed; tarsi not divided into joints; prosternum
small._ The only species of this aberrant family, _Cyathocerus horni_, has
been found in Central America. Nothing is known as to its life-history.
FAM. 46. GEORYSSIDAE.—_Antennae short, clubbed; tarsi four-jointed;
prosternum very small; front coxae exserted, but not contiguous._ There are
about two dozen species of these small beetles known. Our British
_Georyssus pygmaeus_ lives in extremely wet places, and covers itself with
a coating of mud or fine sand so that it can only be detected when in
movement. Nothing further is known as to its life-history or habits.
Members of the genus have been detected in widely-separated parts of the
globe.
FAM. 47. HETEROCERIDAE.—_Labrum and mandibles projecting forwards; antennae
short, the terminal seven joints broad and short, forming a sort of broad
serrate club; legs armed with stout spines; tarsi four-jointed._ The
Heteroceridae are small beetles covered with very dense but minute
pubescence; they live in burrows among mud or sand in wet places, and are
found in many parts of the world. They possess a stridulating organ in the
form of a slightly elevated curved line on each side of the base of the
abdomen, rubbed by the posterior femur. The larvae live in the same places
as the beetles; they have well-developed thoracic legs, the mandibles are
porrect, the three thoracic segments rather large, and the body behind
these becomes gradually narrower; they are believed to eat the mud amongst
which they burrow. We have seven British species of Heteroceridae.
FAM. 48. PARNIDAE.—_Prosternum distinct in front of the coxae, usually
elongate, behind forming a process received into a definite cavity on the
mesosternum; head retractile, the mouth protected by the prosternum. Tarsi
five-jointed, terminal joint long._ {244}Although the characters of these
Insects are not very different from those of Byrrhidae, of Dascillidae, and
even of certain Elateridae, there is practically but little difficulty in
distinguishing Parnidae. They are of aquatic habits, though many, in the
perfect state, frequently desert the waters. There are about 300 or 400
species known, but the family is doubtless more extensive, as these small
beetles attract but little notice. There are two groups:—1. Parnides, in
which the front coxae have a considerable transverse extension, the
antennae are frequently short and of peculiar structure, and the body is
usually clothed with a peculiar, dense pubescence. 2. Elmides, with round
front coxae, a bare, or feebly pubescent body, and simple antennae.
_Parnus_ is a genus commonly met with in Europe, and is less aquatic in
habits than its congeners; it is said to enter the water carrying with it a
coating of air attached to its pubescence. Its larvae are not well known;
they live in damp earth near streams, and are said to much resemble the
larvae of Elateridae. _Potamophilus acuminatus_ has a very interesting
larva, described by Dufour; it lives on decaying wood in the Adour. It is
remarkable from the ocelli being arranged so as to form an almost true eye
on each side of the head; there are eight pairs of abdominal spiracles, and
also a pair on the mesothorax, though there are none on the pro- or
meta-thorax; each of the stigmata has four elongate sacs between it and the
main tracheal tube; the body is terminated by a process from which there
can be protruded bunches of filamentous branchiae. The larvae of
_Macronychus quadrituberculatus_ is somewhat similar, though the features
of its external structure are less remarkable. The Elmides live attached to
stones in streams; the larva is rather broad, fringed at the sides of the
body, and bears behind three elegant sets of fine filamentous branchiae.
The North American genus _Psephenus_ is placed in Parnidae, though instead
of five, the male has seven, the female six, visible ventral segments; the
larva is elliptical, with dilated margins to the body. Friederich, has
given,[123] without mentioning any names, a detailed account of Brazilian
Parnid larvae, that may perhaps be allied to _Psephenus_.
FAM. 49. DERODONTIDAE.—_Tarsi five-jointed, slender, fourth joint rather
small; front coxae prominent and transversely {245}prolonged; middle coxae
small; abdomen with five visible segments, all mobile, the first not
elongated._ One of the smallest and least known of the families of
Coleoptera; it consists of four or five species of small Insects of the
genera _Derodontus_ and _Peltasticta_, found in North America, Europe, and
Japan. The distinction of the family from Cleridae is by no means certain;
our European _Laricobius_ apparently possessing characters but little
different. Nothing is known as to the life-histories.
FAM. 50. CIOIDAE.—_Small or minute beetles; antennae short, terminal joints
thicker; tarsi short, four-jointed; anterior and middle coxae small, oval,
deeply embedded; abdomen with five ventral segments, all mobile._ The
position of these obscure little Insects seems to be near Colydiidae and
Cryptophagidae, though they are usually placed near Bostrichidae. So far as
known, they all live in fungi, or in wood penetrated by fungoid growths.
The cylindrical larvae live also in similar matter; they usually have the
body terminated behind by one or two hooks curved upwards; that of _Cis
melliei_ (Fig. 124) has, instead of these hooks, a curious chitinous tube.
About 300 species of the family are now known; a score, or so, occurring in
Britain. The Hawaiian Islands have a remarkably rich and varied fauna of
Cioidae.
[Illustration: Fig. 123—_Derodontus maculatus._ North America.]
[Illustration: Fig. 124—_Cis melliei._ Martinique. A, Perfect Insect; B,
pupa; C, larva; D, terminal portion of body of larva. (After Coquerel.)]
FAM. 51. SPHINDIDAE.—This family of half a dozen species of rare and small
Insects, differs from Cioidae by the tarsi being five-jointed at any rate
on the front and middle feet, opinions differing as to whether the number
of joints of the hind tarsi is four or five. These Insects live in fungi
growing in wood, _e.g._ _Reticularia hortensis_, that are at first pulpy
and {246}afterwards become powder. The larvae of both of our British
genera, _Sphindus_ and _Aspidiphorus_, have been described by Perris, who
considers them allied to the fungivorous Silphidae and Latridiidae. The
systematic position of these Insects has been the subject of doubt since
the days of Latreille.
FAM. 52. BOSTRICHIDAE (APATIDAE of some authors).—_Tarsi five-jointed, but
the first joint very short and imperfectly separated from the second; front
coxae prominent, contiguous, very little extended transversely; five
visible ventral segments._ The Bostrichidae attack dry wood, and sometimes
in such large numbers that timber is entirely destroyed by them; most of
them make cylindrical burrows into the wood. The larvae have the posterior
part of the body incurved, and resemble the wood-boring larvae of Anobiidae
rather than the predaceous larvae of Cleridae. We follow Leconte and Horn
in placing Lyctides as a division of Bostrichidae; although differing very
much in appearance, they have similar habits and larvae. The typical
Bostrichides are remarkable for their variety of sculpture and for the
shapes of the posterior part of the body; this part is more or less
conspicuously truncate, and furnished with small prominences. _Dinapate
wrightii_, found in the stems of a species of _Yucca_ in the Mojave desert
of California, attains a length of nearly two inches; its larva is
extremely similar to that of _A. capucina_. Some of the forms (_Phonapate_)
stridulate in a manner peculiar to themselves, by rubbing the front leg
against some projections at the hind angle of the prothorax. Upwards of 200
species of the family are known. In Britain we have only four small and
aberrant forms.
[Illustration: Fig. 125—_Apate capucina._ Europe. A, Larva (after Perris);
B, perfect Insect.]
FAM. 53. PTINIDAE.—_Tarsi five-jointed, first joint not reduced in size,
often longer than second; front and middle coxae small, not transversely
extended, the former slightly prominent; five visible ventral segments;
prosternum very short._ Here are included two sub-families, Ptinides and
Anobiides; they are considered as distinct families by many authors, but in
the present imperfect state of knowledge[124] it is not necessary to treat
them separately.
{247}[Illustration: Fig. 126—"Biscuit-weevil." _Anobium paniceum._]
[Illustration: Fig. 127.—Early stages of _Anobium paniceum_. A, Eggs,
variable in form; B, larva; C, pupa; D, asymmetrical processes terminating
body of pupa. [This larva is probably the "book-worm" of librarians].]
Ptinidae are sometimes very destructive to dried animal matter, and attack
specimens in museums; Anobiides bore into wood, and apparently emerge as
perfect Insects only for a very brief period; _Anobium_ (_Sitodrepa_)
_paniceum_ is, however, by no means restricted in its tastes; it must
possess extraordinary powers of digestion, as we have known it to pass
several consecutive generations on a diet of opium; it has also been
reported to thrive on tablets of dried compressed meat; in India it is said
to disintegrate books; a more usual food of the Insect is, however, hard
biscuits; weevilly biscuits are known to every sailor, and the so-called
"weevil" is usually the larva of _A. paniceum_ (Fig. 127, B). In the case
of this Insect we have not detected more than one spiracle (situate on the
first thoracic segment); the other known larvae of Anobiides are said to
possess eight abdominal spiracles. The skeleton in some of this sub-family
is extremely modified, so as to allow the Insects to pack themselves up in
repose; the head is folded in over the chest, and a cavity existing on the
breast is thus closed by the head; in this cavity the antennae and the
prominent mouth-parts are received and protected; the legs shut together
{248}in an equally perfect manner, so that no roughness or chink remains,
and the creature looks like a little hard seed. _Anobium striatum_ is a
common Insect in houses, and makes little round holes in furniture, which
is then said to be "worm-eaten." _A._ (_Xestobium_) _tessellatum_, a much
larger Insect, has proved very destructive to beams in churches, libraries,
etc. These species are the "death-watches" or "greater death-watches" that
have been associated with the most ridiculous superstitions (as we have
mentioned in Volume V., when speaking of the lesser death-watches, or
Psocidae). The ticking of these Insects is really connected with sex, and
is made by striking the head rapidly against the wood on which the Insect
is standing.
The very anomalous genus _Ectrephes_ (Fig. 128) is found in ants' nests in
Australia. Westwood placed it in Ptinidae. Wasmann has recently treated it
as a distinct family, Ectrephidae, associating it with _Polyplocotes_ and
_Diplocotes_, and treating them as allied to Scydmaenidae.
[Illustration: Fig. 128—_Ectrephes kingi._ West Australia. (After
Westwood.)]
FAM. 54. MALACODERMIDAE.—_Seven (or even eight) visible ventral segments,
the basal one not co-adapted in form with the coxae; tarsi five-jointed.
Integument softer than usual, the parts of the body not accurately
co-adapted._ This important family includes a variety of forms: viz.
Lycides, Drilides, Lampyrides, Telephorides; though they are very different
in appearance, classifiers have not yet agreed on separating them as
families. Of these the Lampyrides, or glow-worms, are of special interest,
as most of their members give off a phosphorescent light when alive; in
many of them the female is apterous and like a larva, and then the light it
gives is usually conspicuous, frequently much more so than that of its
mate; in other cases the males are the most brilliant. The exact importance
of these characters in the creatures' lives is not yet clear, but it
appears probable that in the first class of cases the light of the female
serves as an attraction to the male, while in the second class the very
brilliant lights of the male serve as an amusement, or as an incitement to
rivalry amongst the individuals of this sex.
{249}[Illustration: Fig. 129—_Phengodes hieronymi._ Cordoba, South America.
(After Haase.) A, Male; B, female. _l_, _l_, Positions of luminous spots;
_ls_, spiracles. About × 3.]
The well-known fire-flies (_Luciola_) of Southern Europe are an example of
the latter condition. They are gregarious, and on calm, warm nights crowds
of them may be seen moving and sparkling in a charming manner. These
individuals are all, or nearly all, males; so rare indeed is the female
that few entomologists have even noticed it. The writer once assisted in a
large gathering of _Luciola italica_ in the Val Anzasca, which consisted of
many hundreds of specimens; all of those he caught, either on the wing or
displaying their lights on the bushes, were males, but he found a solitary
female on the ground. This sex possesses ordinary, small eyes instead of
the large, convex organs of the male, and its antennae and legs are much
more feeble, so that though provided with elytra and wings it is altogether
a more imperfect creature. Emery has given an account of his observations
and experiments on this Insect, but they do not give any clear idea as to
the exact function of the light.[125] In our British glow-worm the female
is entirely apterous—hence the name glow-worm—but the male has elytra and
ample wings, and frequently flies at night into lighted apartments.
Although so little has been ascertained as to the light of Lampyridae,
there are two facts that justify us in supposing that it is in some way of
importance to the species. These are: (1) that in a great many species the
eyes have a magnificent and unusual development; (2) that the habits of the
creatures are in nearly all cases nocturnal. It is true that the little
_Phosphaenus hemipterus_ is said to be diurnal in habits, but it is
altogether an exceptional form, being destitute of wings in both sexes, and
possessed of only very feeble light-giving powers, and we have, moreover,
very little real knowledge as to its natural history; it is said {250}that
the female is of the utmost rarity, though the male is not uncommon.
The nature of the luminosity of _Lampyris_ has given rise to many
contradictory statements; the light looks somewhat like that given off by
phosphorus, and is frequently spoken of as phosphorescence; but
luminescence is a better term. The egg, larva, pupa, and male are luminous
as well as the female (at any rate in _L. noctiluca_); the luminescence is,
however, most marked in the female imago, in which it is concentrated near
the extremity of the abdomen; here there are two strata of cells, and many
fine capillary tracheae are scattered through the luminous substance.
Wielowiejski concludes that the light-producing power is inherent in the
cells of the luminous organ, and is produced by the slow oxidation of a
substance formed under the influence of the nervous system. The cells are
considered to be essentially similar to those of the fat-body.[126] The
luminescence of Lampyridae is very intermittent, that is to say, it is
subject to rapid diminutions and increases of its brilliancy; various
reasons have been assigned for this, but all are guesses, and all that can
be said is that the changes are possibly due to diminution or increase of
the air-supply in the luminous organ, but of the way in which this is
controlled there seems to be no evidence. Considerable difference of
opinion has existed as to the luminescence of the eggs of _Lampyris_. If it
exist in the matter contained in the egg, it is evident that it is
independent of the existence of tracheae or of a nervous system. Newport
and others believed that the light given by the egg depended merely on
matter on its exterior. The observations of Dubois[127] show, however, that
it exists in the matter in the egg; he has even found it in the interior of
eggs that had been deposited unfertilised.
From time to time, since the commencement of the nineteenth century, there
have appeared imperfect accounts of extraordinary light-giving larvae found
in South America, of various sizes, but attaining in some cases a length,
it is said, of three inches; they are reported as giving a strong red light
from the two extremities of the body, and a green light from numerous
points along the {251}sides of the body, and hence are called, it is said,
in Paraguay the railway-beetle. We may refer the reader to Haase's
paper[128] on the subject of these "larvae," as we can here only say that
it appears probable that most of these creatures may prove to be adult
females of the extraordinary group Phengodini, in which it would appear
that the imago of the female sex is in a more larva-like state than it is
in any other Insects. The males, however, are well-developed beetles;
unlike the males of Lampyrides, in general they have not peculiar eyes, but
on the other hand they possess antennae which are amongst the most highly
developed known, the joints being furnished on each side with a long
appendage densely covered with pubescence of a remarkable character. There
is no reason to doubt that Haase was correct in treating the Insect we
figure (Fig. 129, B) as a perfect Insect; he is, indeed, corroborated by
Riley.[129] The distinctions between the larva and female imago are that
the latter has two claws on the feet instead of one, a greater number of
joints in the antennae, and less imperfect eyes; the female is in fact a
larva, making a slightly greater change at the last ecdysis, than at those
previous. It is much to be regretted that we have so very small a knowledge
of these most interesting Insects. Malacodermidae are probably the most
imperfect or primitive of all beetles, and it is a point of some interest
to find that in one of them the phenomena of metamorphosis are reduced in
one sex to a minimum, while in the other they are—presumably at
least—normal in character.
Numerous larvae of most extraordinary, though diverse, shapes, bearing long
processes at the sides of the body, and having a head capable of complete
withdrawal into a slender cavity of the thorax, have long been known in
several parts of the world, and Dr. Willey recently found in New Britain a
species having these body-processes articulated. Though they are doubtless
larvae of Lampyrides, none of them have ever been reared or exactly
identified.
A very remarkable Ceylonese Insect, _Dioptoma adamsi_ Pascoe, is placed in
Lampyrides, but can scarcely belong there, as apparently it has but five or
six visible ventral segments; this Insect has two pairs of eyes, a large
pair, with coarse facets on {252}the under side of the head, and a
moderate-sized pair with fine facets on the upper side. Nothing is known as
to the habits of this curiosity, not even whether it is luminous in one or
both sexes.
It is believed that the perfect instar of Lampyrides takes no food at all.
The larvae were formerly supposed to be vegetarian, but it appears probable
that nearly all are carnivorous, the chief food being Mollusca either
living or dead. The larvae are active, and in many species look almost as
much like perfect Insects as do the imagos.
The other divisions of Malacodermidae—Lycides, Drilides, Telephorides—also
have predaceous, carnivorous larvae. All these groups are extensive. Though
much neglected by collectors and naturalists, some 1500 species of the
family Malacodermidae have been detected. We have about 50 in Britain, and
many of them are amongst the most widely distributed and abundant of our
native Insects. Thus, however near they may be to the primitive condition
of Coleoptera, it is highly probable that they will continue to exist
alongside of the primitive Cockroaches and Aptera, long after the more
highly endowed forms of Insect-life have been extinguished wholesale by the
operations of mankind on the face of the earth.
[Illustration: Fig. 130—_Malachius aeneus._ Britain. A, Larva (after
Perris); B, female imago.]
FAM. 55. MELYRIDAE (or MALACHIIDAE).—_Six visible and moveable ventral
abdominal segments; the basal part more or less distinctly co-adapted with
the coxae._ These Insects are extremely numerous, but have been very little
studied. In many works they are classified with Malacodermidae, but were
correctly separated by Leconte and Horn, and this view is also taken by Dr.
Verhoeff, the latest investigator. The smaller number of visible ventral
segments appears to be due to a change at the base correlative with an
adaptation between the base of the abdomen and the hind coxae. The
characters are singularly parallel with those of Silphidae; but in
Melyridae the antennae are filiform or serrate, {253}not clavate. The
habits in the two families are different, as the Melyridae are frequenters
of flowers. Many of the Melyridae have the integument soft, but in the
forms placed at the end of the family—_e.g._ _Zygia_—they are much firmer.
Thus these Insects establish a transition from the Malacodermidae to
ordinary Coleoptera. Although the imagos are believed to consume some
products of the flowers they frequent, yet very little is really known, and
it is not improbable that they are to some extent carnivorous. This is the
case with the larvae that are known (Fig. 130, larva of _Malachius
aeneus_). These are said by Perris to bear a great resemblance to those of
the genus _Telephorus_, belonging to the Malacodermidae.
FAM. 56. CLERIDAE.—_Tarsi five-jointed; but the basal joint of the
posterior very indistinct, usually very small above, and closely united
with the second by an oblique splice; the apices of joints two to four
usually prolonged as membranous flaps; anterior coxae prominent, usually
contiguous, rather large, but their cavities not prolonged externally;
labial palpi usually with large hatchet-shaped terminal joint; ventral
segments five or six, very mobile._ The Cleridae are very varied in form
and colours; the antennae are usually more or less clubbed at the tip, and
not at all serrate, but in _Cylidrus_ and a few others they are not
clubbed, and in _Cylidrus_ have seven flattened joints. The student should
be very cautious in deciding as to the number of joints in the feet in this
family, as the small basal joint is often scarcely distinguishable, owing
to the obliteration of its suture with the second joint. The little Alpine
_Laricobius_ has the anterior coxal cavities prolonged externally, and the
coxae receive the femora to some extent, so that it connects Cleridae and
Derodontidae. The Cleridae are predaceous, and their larvae are very
active; they are specially fond of wood-boring Insects; that of _Tillus
elongatus_ (Fig. 131) enters the burrows of _Ptilinus pectinicornis_ in
search of the larva. The members of the group Corynetides frequent animal
matter, carcases, bones, etc., and, it is said, feed thereon, but Perris's
recent investigations[130] make it probable that the larvae really eat the
innumerable Dipterous larvae found in such refuse; it is also said that the
larvae of Cleridae spin cocoons for their metamorphosis; but Perris has
also shown that the larvae of _Necrobia ruficollis_ really use the puparia
formed {254}by Diptera. Some of the species of _Necrobia_ have been spread
by commercial intercourse, and _N. rufipes_ appears to be now one of the
most cosmopolitan of Insects. The beautifully coloured _Corynetes
coeruleus_ is often found in our houses, and is useful, as it destroys the
death-watches (_Anobium_) that are sometimes very injurious. _Trichodes
apiarius_, a very lively-coloured red and blue beetle, destroys the larvae
of the honey-bee, and Lampert has reared _Trichodes alvearius_ from the
nests of _Chalicodoma muraria_, a mason-bee; he records that one of its
larvae, after being full grown, remained twenty-two months quiescent and
then transformed to a pupa. Still more remarkable is a case of fasting of
the larva of _Trichodes ammios_ recorded by Mayet;[131] this Insect, in its
immature form, destroys _Acridium maroccanum_; a larva sent from Algeria to
M. Mayet refused such food as was offered to it for a period of two and a
half years, and then accepted mutton and beef as food; after being fed for
about a year and a half thereon, it died. Some Cleridae bear a great
resemblance to Insects of other families, and it appears probable that they
resemble in one or more points the Insects on which they feed. The species
are now very numerous, about 1000 being known, but they are rare in
collections; in Britain we have only nine species, and some of them are now
scarcely ever met with.
[Illustration: Fig. 131—Larva of _Tillus elongatus_. (New Forest). A, Head;
B, front leg; C, termination of the body, more magnified.]
FAM. 57. LYMEXYLONIDAE.—_Elongate beetles, with soft integuments, front and
middle coxae exserted, longitudinal in position; tarsi slender,
five-jointed; antennae short, serrate, but rather broad._ Although there
are only twenty or thirty species of this family, they occur in most parts
of the world, and are remarkable on account of their habit of drilling
cylindrical holes in hard wood, after the manner of Anobiidae. The larva of
_Lymexylon navale_ was formerly very injurious to timber used for
constructing ships, but of late years its ravages appear to have been of
little importance. The genus _Atractocerus_ consists of a few species of
very abnormal {255}Coleoptera, the body being elongate and vermiform, the
elytra reduced to small, functionless appendages, while the wings are
ample, not folded, but traversed by strong longitudinal nervures, and with
only one or two transverse nervures. Owing to the destruction of our
forests the two British Lymexylonidae—_L. navale_ and _Hylecoetus
dermestoides_—are now very rarely met with.
[Illustration: Fig. 132—_Hydrocyphon deflexicollis._ Britain. A, Larva
(after Tournier); B, imago.]
FAM. 58. DASCILLIDAE.—_Small or moderate-sized beetles, with rather flimsy
integuments, antennae either serrate, filiform, or even made flabellate by
long appendages; front coxae elongate, greatly exserted; abdomen with five
mobile ventral segments; tarsi five-jointed._ This is one of the most
neglected and least known of all the families of Coleoptera, and one of the
most difficult to classify; though always placed amongst the Serricornia,
it is more nearly allied to Parnidae and Byrrhidae, that are placed in
Clavicornia, than it is to any of the ordinary families of Serricornia. It
is probable that careful study will show that it is not natural as at
present constituted, and that the old families, Dascillidae and Cyphonidae,
now comprised in it, will have to be separated. Only about 400 species are
at present known; but as nearly 100 of these have been detected in New
Zealand, and 17 in Britain, doubtless the numbers in other parts of the
world will prove very considerable, these Insects having been neglected on
account of their unattractive exterior, and fragile structure. The few
larvae known are of three or four kinds. That of _Dascillus cervinus_ is
subterranean, and is believed to live on roots; in form it is somewhat like
a Lamellicorn larva, but is straight, and has a large head. Those of the
Cyphonides are aquatic, and are remarkable for possessing antennae
consisting of a great many joints (Fig. 132, A). Tournier describes the
larva of _Helodes_ as possessing abdominal but not thoracic spiracles, and
as breathing by coming to the surface of the water and carrying down a
bubble of air adhering to the posterior part of the body; the larva of
_Hydrocyphon_ (Fig. 132, A) {256}possesses several finger-like pouches that
can be exstulpated at the end of the body. It is probable that these larvae
are carnivorous. The imago of this Insect abounds on the bushes along the
banks of some of the rapid waters of Scotland; according to Tournier, when
alarmed, it enters the water and goes beneath it for shelter. The third
form of larva belongs to the genus _Eucinetus_, it lives on fungoid matter
on wood, and has ordinary antennae of only four joints.[132] It is very
doubtful whether _Eucinetus_ is related to other Dascillidae; some
authorities indeed place it in Silphidae.
FAM. 59. RHIPICERIDAE.—_Tarsi five-jointed, furnished with a robust
onychium (a straight chitinous process bearing hairs) between the claws;
antennae of the male bearing long processes, and sometimes consisting of a
large number of joints. Mandibles robust, strongly curved, and almost
calliper-like in form._ This small family of less than 100 species is
widely distributed, though confined to the warmer regions of the earth, a
single species occurring in the extreme south of Eastern Europe. Very
little is known as to the natural history. The larva of _Callirhipis
dejeani_ (Fig. 133, A) is described by Schiödte as hard, cylindrical in
form, and peculiarly truncate behind, so that there appear to be only eight
abdominal segments, the ninth segment being so short as to look like an
operculum at the extremity of the body. It lives in wood.
[Illustration: Fig. 133—A, Larva of _Callirhipis dejeani_ (after Schiödte);
B, _Rhipicera mystacina_ male, Australia; C, under side of its hind foot.]
{257}[Illustration: Fig. 134—_Athous rhombeus._ New Forest. A, Larva; B,
female imago.]
FAM. 60. ELATERIDAE (_Click-beetles_).—_Antennae more or less serrate along
the inner margin, frequently pectinate, rarely filiform. Front coxae small,
spherical. Thorax usually with hind angles more or less prolonged
backwards; with a prosternal process that can be received in, and usually
can move in, a mesosternal cavity. Hind coxa with a plate, above which the
femur can be received. Visible ventral segments usually five, only the
terminal one being mobile. Tarsi five-jointed._ This large family of
Coleoptera comprises about 7000 species. Most of them are readily known by
their peculiar shape, and by their faculty of resting on the back,
stretching themselves out flat, and then suddenly going off with a click,
and thus jerking themselves into the air. Some, however, do not possess
this faculty, and certain of these are extremely difficult to recognise
from a definition of the family. According to Bertkau[133] our British
_Lacon murinus_ is provided near the tip of the upper side of the abdomen
with a pair of eversible glands, comparable with those that are better
known in Lepidopterous larvae. He states that this Insect does not try to
escape by leaping, but shams death and "stinks away" its enemy. The glands,
it would appear, become exhausted after the operation has been repeated
many times. The extent of the leap executed by click-beetles differs
greatly; in some species it is very slight, and only just sufficient to
turn the Insect right side up when it has been placed on its back. In some
cases the Insects go through the clicking movements with little or no
appreciable result in the way of consequent propulsion. Although it is
difficult to look on this clicking power as of very great value to the
Elateridae, yet their organisation is profoundly modified so as to permit
its accomplishment. The junction of the prothorax with the after-body
involves a large number of pieces which are all more or less changed, so
that the joint is endowed with greater mobility than usual; while in the
position of repose, on the other hand, the two parts are firmly locked
together. The thoracic stigma is of a highly remarkable nature, and the
extensive {258}membrane in which it is placed appears to be elastic.
Although the mechanics of the act of leaping are still obscure, yet certain
points are clear; the prosternal process possesses a projection, or notch,
on its upper surface near the tip; as a preliminary to leaping, this
projection catches against the edge of the mesosternal cavity, and as long
as this position is maintained the Insect is quiescent; suddenly, however,
the projection slips over the catch, and the prosternal process is driven
with force and rapidity into the mesosternal cavity pressing against the
front wall thereof, and so giving rise to the leap.
Several larvae are well known; indeed the "wire-worms" that are sometimes
so abundant in cultivated places are larvae of Elateridae. In this instar
the form is usually elongate and nearly cylindrical; the thoracic segments
differ but little from the others except that they bear rather short legs;
the skin is rather hard, and usually bears punctuation or sculpture; the
body frequently terminates in a very hard process, of irregular shape and
bearing peculiar sculpture on its upper surface, while beneath it the
prominent anal orifice is placed: this is sometimes furnished with hooks,
the function of which has not yet been observed. The majority of these
larvae live in decaying wood, but some are found in the earth; as a rule
the growth is extremely slow, and the life of the larva may extend over two
or more years. Some obscurity has prevailed as to their food; it is now
considered to be chiefly flesh, though some species probably attack
decaying roots; and it is understood that wire-worms destroy the living
roots, or underground stems, of the crops they damage. Various kinds of
Myriapods (see Vol. V. p. 29) are often called "wire-worm," but they may be
recognised by possessing more than six legs. The larvae of the genus
_Cardiophorus_ are very different, being remarkably elongate without the
peculiar terminal structure, but apparently composed of twenty-three
segments.
The genus _Pyrophorus_ includes some of the most remarkable of light-giving
Insects. There are upwards of 100 species, exhibiting much diversity as to
the luminous organs; some are not luminous at all; but all are peculiar to
the New World, with the exception that there may possibly be luminous
species, allied to the American forms, in the Fiji Islands and the New
Hebrides. In the tropics of America the _Pyrophorus_, or Cucujos, form one
of the most remarkable of the natural phenomena. {259}The earliest European
travellers in the New World were so impressed by these Insects that
descriptions of their wondrous display occupy a prominent position in the
accounts of writers like Oviedo, whose works are nearly 400 years old. Only
one of the species has, however, been investigated. _P. noctilucus_ is one
of the most abundant and largest of the _Pyrophorus_, and possesses on each
side of the thorax a round polished space from which light is given forth;
these are the organs called eyes by the older writers. Besides these two
eye-like lamps the Insect possesses a third source of light situate at the
base of the ventral surface of the abdomen; there is no trace of this
latter lamp when the Insect is in repose; but when on the wing the abdomen
is bent away from the breast, and then this source of light is exposed;
hence, when flying, this central luminous body can be alternately displayed
and concealed by means of slight movements of the abdomen. The young larva
of _P. noctilucus_ is luminous, having a light-giving centre at the
junction of the head and thorax; the older larva has also numerous luminous
points along the sides of the body near the spiracles. It is remarkable
that there should be three successive seats of luminescence in the life of
the same individual. The eggs too are said to be luminous. The light given
off by these Insects is extremely pleasing, and is used by the natives on
nocturnal excursions, and by the women for ornament. The structure of the
light-organs is essentially similar to that of the Lampyridae. The light is
said to be the most economical known; all the energy that is used being
converted into light, without any waste by the formation of heat or
chemical rays. The subject has been investigated by Dubois,[134] who comes,
however, to conclusions as to the physiology of the luminous processes
different from those that have been reached by Wielowiejski and others in
their investigations on Glow-worms. He considers that the light is produced
by the reactions of two special substances, luciferase and luciferine.
Luciferase is of the nature of an enzyme, and exists only in the luminous
organs, in the form, it is supposed, of extremely minute granules.
Luciferine exists in the blood; and the light is actually evoked by the
entry of blood into the luminous organ.
We have given to this family the extension assigned to it by {260}Schiödte.
Leconte and Horn also adopt this view, except that they treat Throscides as
a distinct family. By most authors Eucnemides, Throscides, and Cebrionides
are all considered distinct families, but at present it is almost
impossible to separate them on satisfactory lines. The following table from
Leconte and Horn exhibits the characters of the divisions so far as the
imago is concerned:—
Posterior coxae laminate; trochanters small.
Labrum concealed; antennae somewhat distant from the eyes,
their insertion narrowing the front Eucnemides.
Labrum visible, free; antennae arising near the eyes under
the frontal margin Elaterides.
Labrum transverse, connate with the front.
Ventral segments six; claws simple; tibial spurs well
developed. Cebrionides.
Ventral segments five; claws serrate; tibial spurs moderate.
Perothopides.
Posterior coxae not laminate; trochanters of middle and
posterior legs very long Cerophytides.
[Illustration: Fig. 135—Larva of _Fornax_ n. sp. Hawaii. A, Upper side; B,
under side: _s_ _s_, position of spiracles; C, head more enlarged; D, under
side of terminal segment; _a_, anus.]
Throscides are considered to be distinguished by the mesosternum being
impressed on each side in front for the accommodation of the posterior face
of the front coxae. The genus _Throscus_ has the antennae clavate. The
classification of the Elaterides and these forms is a matter of the
greatest difficulty, and, if the larvae are also considered, becomes even
more complex. Cebrionid larvae are different from those of any of the other
divisions, and possess laminate, not calliper-like, mandibles. The larvae
of Eucnemides (Fig. 135) are very little known, but are highly remarkable,
inasmuch as it is very difficult to find any mouth-opening in some of them,
and they have no legs. The other divisions possess very few species
compared with Elaterides. In Britain we have about sixty species of
Elaterides, four of Throscides and three of Eucnemides; _Cerophytum_ was
probably a native many years ago. Neither Perothopides {261}nor Cebrionides
are represented in our fauna; the former of these two groups consists only
of four or five North American species, and the Cerophytides are scarcely
more numerous.
FAM. 61. BUPRESTIDAE.—_Antennae serrate, never elongate; prothorax fitting
closely to the after-body, with a process received into a cavity of the
mesosternum so as to permit of no movements of nutation. Five visible
ventral segments, the first usually elongate, closely united with the
second, the others mobile. Tarsi five-jointed, the first four joints
usually with membranous pads beneath._ This family is also of large extent,
about 5000 species being known. Many of them are remarkable for the
magnificence of their colour, which is usually metallic, and often of the
greatest brilliancy; hence their wing-cases are used by our own species for
adornment. The elytra of the eastern kinds of the genus _Sternocera_ are of
a very brilliant green colour, and are used extensively as embroidery for
the dresses of ladies; the bronze elytra of _Buprestis (Euchroma) gigantea_
were used by the native chieftains in South America as leg-ornaments, a
large number being strung so as to form a circlet. The integument of the
Buprestidae is very thick and hard, so as to increase the resemblance to
metal. The dorsal plates of the abdomen are usually soft and colourless in
beetles, but in Buprestidae they are often extremely brilliant. The
metallic colour in these Insects is not due to pigment, but to the nature
of the surface. Buprestidae appear to enjoy the hottest sunshine, and are
found only where there is much summer heat. Australia and Madagascar are
very rich in species and in remarkable forms of the family, while in
Britain we possess only ten species, all of which are of small size, and
nearly all are excessively rare. The family is remarkably rich in fossil
forms; no less than 28 per cent of the Mesozoic beetles found by Heer in
Switzerland are referred to Buprestidae.
[Illustration: Fig. 136—A, Larva of _Euchroma goliath_ (after Schiödte); B,
imago of _Melanophila decostigma_. Europe.]
The larvae (Fig. 136, A) find nourishment in living vegetable matter, the
rule being that they form galleries in or under the {262}bark of trees and
bushes, or in roots thereof; some inhabit the stems of herbaceous plants
and one or two of the smaller forms have been discovered to live in the
parenchyma of leaves. A few are said to inhabit dead wood, and in Australia
species of _Ethon_ dwell in galls on various plants. Buprestid larvae are
of very remarkable shape, the small head being almost entirely withdrawn
into the very broad thorax, while the abdomen is slender.[135] A few,
however, depart from this shape, and have the thoracic region but little or
not at all broader than the other parts. The larvae of _Julodis_—a genus
that inhabits desert or arid regions—are covered with hair; they have a
great development of the mandibles; it is believed that they are of
subterranean habits, and that the mandibles are used for burrowing in the
earth. Only the newly hatched larva is, however, known.
SERIES IV. HETEROMERA.
_Tarsi of the front and middle legs with five, those of the hind legs with
four, joints._
This series consists of some 14,000 or 15,000 species. Twelve or more
families are recognised in it, but the majority of the species are placed
in the one great family, Tenebrionidae. The number of visible ventral
segments is nearly always five. Several of the families of the series are
of doubtful validity; indeed beyond that of Tenebrionidae the taxonomy of
this series is scarcely more than a convention. The larvae may be
considered as belonging to three classes; one in which the body is
cylindrical and smooth and the integument harder than usual in larvae; a
second in which it is softer, and frequently possesses more or less
distinct pseudopods, in addition to the six thoracic legs; and a third
group in which hypermetamorphosis prevails, the young larvae being the
creatures long known as Triungulins, and living temporarily on the bodies
of other Insects, so that they were formerly supposed to be parasites.
{263}FAM. 62. TENEBRIONIDAE.—_Front coxae short, not projecting from the
cavities, enclosed behind. Feet destitute of lobed joints. Claws smooth._
This is one of the largest families of Coleoptera, about 10,000 species
being already known. A very large portion of the Tenebrionidae are entirely
terrestrial, wings suitable for flight being absent, and the elytra
frequently more or less soldered. Such forms are described in systematic
works as apterous. Unfortunately no comprehensive study has ever been made
of the wings or their rudiments in these "apterous forms."[136] it is
probable that the wings, or their rudiments or vestiges, always exist, but
in various degrees of development according to the species, and that they
are never used by the great majority of the terrestrial forms. Many of the
wood-feeding Tenebrionidae, and the genera usually placed at the end of the
family, possess wings well adapted for flight. The apterous forms are
chiefly ground-beetles, living in dry places; they are very numerous in
Africa, California, and North Mexico. Their colour is nearly always black,
and this is probably of some physiological importance; the integuments are
thick and hard, and if the wing-cases are taken off, it will be found that
they are usually more or less yellow on the inner face, even when jet-black
externally; the external skeleton is very closely fitted together, the
parts that are covered consisting of very delicate membrane; the transition
between the hard and the membranous portions of the external skeleton is
remarkably abrupt. These ground-Tenebrionidae form a very interesting
study, though, on account of their unattractive appearance, they have not
received the attention they deserve.
[Illustration: Fig. 137—_Tenebrio molitor._ Europe, etc. A, Larva
(meal-worm); B, pupa (after Schiödte); C, imago.]
Many of the Tenebrionidae, notwithstanding their dark {264}colours, are
diurnal in habits, and some of them run with extreme velocity in places so
bare and desert that the means of existence of the Insects is a mystery.
Most of the Tenebrionidae, however, shun the light. The food is usually
vegetable matter, and it is apparently preferred in a very dry state. Mr.
Gahan has recently recorded that in _Praogena_ the under surface of the
head has the gular region striate for stridulating purposes. This is the
only instance known of a voice-organ in this situation, and moreover is the
only case in all the Tenebrionidae in which any sound-producing organ has
been discovered. The larvae exhibit but little variety, they are elongate
and cylindrical, with harder integument than is usual in Coleopterous
larvae; they have six thoracic legs, and at the under side of the posterior
extremity the anus serves as a very short pseudopod. The resemblance of
these larvae to those of Elateridae is considerable; but though the body is
terminated by one or two small processes, these never attain the complexity
of the terminal segment of Elateridae. The common meal-worm—_i.e._ the
larva of _Tenebrio molitor_—is a very characteristic example of the group.
The pupae are remarkable on account of peculiar projections, of varied and
irregular form, that exist on the sides of the abdominal segments. Britain
is very poor in these Insects; our list of them scarcely attains the number
of thirty species.
FAM. 63. CISTELIDAE.—_Claws comb-like._ The very obscure beetles forming
this family are only separated from Tenebrionidae on account of their
pectinate claws. About 500 species of Cistelidae are recorded; the early
instars, so far as known, do not differ from those of Tenebrionidae; the
larvae are believed to live on dead wood.
FAM. 64. LAGRIIDAE.—_Anterior coxal cavities closed, tips of the front
coxae free, claws smooth, penultimate joint of the tarsi broader, pubescent
beneath._ This family has very little to distinguish it from Tenebrionidae,
and the group Heterotarsini appears to connect the two. It is a small
family of about 200 species, widely distributed, and represented in Britain
by one species, _Lagria hirta_. The early instars are similar to those of
the Tenebrionidae, except that the larva is less retiring in its habits and
wanders about on foliage: it is of broader form than that of most of the
Tenebrionidae. The pupa has long projections at the sides of the abdominal
segments.
{265}FAM. 65. OTHNIIDAE.—Only about ten species are known of this dubious
family. They are small Insects with weak integument, and are said by
Leconte and Horn to be distinguished from "degraded Tenebrionidae" by the
more mobile abdominal segments, the hind-margins of which are
semi-membranous. The antennae are of the clubbed shape, characteristic of
"Clavicornia," but this also occurs in numerous undoubted Tenebrionidae.
Species of _Othnius_ have been found in Japan and Borneo, as well as in
North America. Nothing is known as to their metamorphoses.
FAM. 66. ÆGIALITIDAE.—_All the coxae very widely separated; no
co-adaptation between the sides of the abdomen and the edges of the
wing-cases; five ventral segments and tip of a sixth visible._ Two minute
and rare Insects from North-West America constitute this family. It is
distinguished from Pythidae by the minute front coxae, widely separated,
completely closed in, and deeply embedded in the prosternum.
FAM. 67. MONOMMIDAE.—This is a small family of less than 100 species, the
members of which have the details of their external structure much
modified, permitting the Insect to pack itself up in repose in a very
perfect manner. They are of small size and oval form; and are absent from
Europe and the Antipodes. Nothing appears to be known as to the
metamorphosis.
FAM. 68. NILIONIDAE.—_Broad, circular Heteromera, of moderate size, with
the front coxae but little separated, and the anterior acetabula closed,
though having the appearance of being open in consequence of the tips of
the epimera being free. The inflexed portion of the wing-cases remarkably
broad._ A small family of less than fifty species, found on fungi, chiefly
in South America. The metamorphoses are not known. It is of very doubtful
validity.
FAM. 69. MELANDRYIDAE.—_Head not constricted behind the eyes; anterior
acetabula not closed; claws smooth. Prothorax broad behind._ These are
loosely-fitted-together Insects, of moderate or small size, frequenting dry
wood or fungi. About 200 species are known, found chiefly in temperate
regions. The few described larvae are rather varied in their details and
cannot be generalised at present. The characters of the members of this
family require fresh investigation.
FAM. 70. PYTHIDAE.—Distinguished from Melandryidae by the {266}prothorax
being narrow behind. This is a small family of about 100 species, found in
temperate regions in connection with timber. The species of _Rhinosimus_
have the head prolonged in front of the antennae so as to form a beak. The
larva of _Pytho depressus_ is flat and has parallel sides; the body is
terminated by two widely-separated sharp processes. It is found
occasionally under the bark of firs in Scotland.
FAM. 71. PYROCHROIDAE.—Differs from Melandryidae by the head forming a very
narrow neck behind, and by the penultimate tarsal joints being broad. They
are feeble Insects, though active on the wing. They are destitute of any of
the various remarkable structures found in Mordellidae. Only about forty
species are known, and the family is confined to the north temperate
region, being best represented in Japan. _Pyrochroa rubens_ is common in
some parts of England; the larva is found under the bark of tree-stumps; it
is remarkably flat, and has the eighth abdominal segment unusually long,
while the ninth terminates the body in the form of two long sharp
processes.
FAM. 72. ANTHICIDAE.—_Head with an abrupt narrow neck; prothorax narrower
than the elytra. Middle and hind coxae placed in definite acetabula. Claws
simple._ These little Insects are numerous in species; they have little
resemblance to Pyrochroidae, though the characters of the two families
cause us to place them in proximity. There are about 1000 species known;
though we have only about 12 in Britain, they are very numerous in the
Mediterranean region. The family Pedilidae of Lacordaire and some others is
now merged in Anthicidae. Thomson and Champion, on the other hand, separate
some very minute Insects to form the family Xylophilidae, on account of
certain differences in the form of the abdomen and tarsi. The Xylophilidae
live in dead wood; the Anthicidae, on the surface of the earth, after the
manner of ground-beetles; very little is, however, known as to their
natural history.
FAM. 73. OEDEMERIDAE.—_Prothorax not forming sharp edges at the sides, head
without a narrow neck. Penultimate tarsal joint broad; claws smooth._ These
Insects usually have a feeble integument, and bear a certain resemblance to
Malacodermidae. Less than 500 species are known, but they are widely
distributed, and occur in both temperate and tropical regions. The larvae
live in old wood. _Nacerdes melanura_ is common on our {267}coasts, where
its larva lives in timber cast up by the sea, or brought down by floods,
and it is able to resist immersion by the tide. It is remarkable from the
possession of five pairs of dorsal false feet on the anterior segments, and
two pairs on the ventral aspect. In _Asclera caerulea_ there are six dorsal
and three ventral pairs of these remarkable pseudopods. We have six species
of Oedemeridae in Britain, including _Asclera_ as well as _Nacerdes_.
[Illustration: Fig. 138—_Asclera caerulea._ A, Larva; B, pupa (after
Schiödte); C, imago. Cambridge.]
FAM. 74. MORDELLIDAE (incl. RHIPIPHORIDAE).—_Head peculiarly formed, vertex
lobed or ridged behind, so that in extension it reposes on the front edge
of the pronotum; capable of great inflection and then covering the
prosternum; hind coxae with laminae forming a sharp edge behind, frequently
very large._ This family is a very distinct one, though it exhibits great
variety. Lacordaire has pointed out that Rhipiphoridae cannot at present be
satisfactorily distinguished from Mordellidae. Leconte and Horn separate
the two by the fact that the sides of the prothorax form a sharp edge in
Mordellidae, but not in Rhipiphoridae. A better character would perhaps be
found by a study of the head, but as this would clearly result in a radical
change in the composition of the two families it is preferable to treat
them at present as only sub-families: if placed on a similar basis to the
preceding families, the group would however form, not two, but several
families. Besides the unusual shape of the head (Fig. 139, D) the ventral
region of the body is remarkably formed, being very convex, and in many
Mordellides terminating in a strong spinous process (Fig. 139, C). The
elytra are, in several Rhipiphorids, of the groups Myoditini and
Rhipidiini, reduced to a very small size, and the wings are not folded. The
Mordellidae are remarkable for their activity; in the perfect state they
usually frequent flowers, and fly and run with extreme rapidity.
Mordellides are amongst the most numerous and abundant of the European
Coleoptera, and in Britain the Anaspini swarm on the flowers of bushes and
Umbelliferae. The {268}life-histories appear to be singularly varied; but
unfortunately they are incompletely known. The larvae of some of the
Mordellids have been found in the stems of plants, and derive their
nutriment therefrom. This is said by Schwarz to be undoubtedly the case
with _Mordellistena floridensis_. Coquillett has found the larvae of _M.
pustulata_ in plant-stems under circumstances that render it highly
probable that they were feeding on a Lepidopterous larva contained in the
stems; and Osborn found a similar larva that was pretty certainly a
_Mordellistena_, and fed voraciously on Dipterous larvae in the stems of a
plant. The little that is known as to the metamorphoses of _Mordella_ and
_Anaspis_ shows that they live in old wood, but does not make clear the
nature of their food.
[Illustration: Fig. 139—_Mordellistena floridensis._ America. (After
Riley.) A, Larva; B, pupa; C, imago; D, outline of detached head of imago
of _M. pumila_, to show the neck.]
Although it has been ascertained that the Rhipiphorides exhibit instances
of remarkable metamorphosis, their life-histories are still very
imperfectly known. Dr. Chapman has ascertained some particulars as to
_Metoecus paradoxus_, which has long been known to prey in the larval state
on the larvae of the common social wasps.[137] The eggs are apparently not
deposited in the nests of the wasps, but in old wood. The young larva is a
triungulin, similar to that of the Cantharidae, we shall subsequently
describe. It is not known how it makes its way to the wasps' nests, but it
is possible that when a wasp visits some old wood haunted by these larvae,
some of them may attach themselves to it and be carried to the wasps'
nests. When {269}access is gained to the cells the little _Metoecus_
pierces the skin of one of the wasp-grubs, and entering in it feeds on the
interior; after it has increased in size it emerges, changes its skin, and
assumes a different form and habits; subsequently, as an external parasite,
entirely devouring the wasp-larva, and then becoming a pupa, and finally a
perfect _Metoecus_, in the cell of the wasp. The wasps, though they
investigate the cells, do not apparently entertain any objection to the
_Metoecus_, though there may be sometimes as many as twenty or thirty of
the destroyers in a single nest. A few hours after the _Metoecus_ has
become a winged Insect and has escaped from the cells, it appears however,
from the observations of Erné[138] on nests of wasps in captivity, that the
wasps become hostile to the foreigners, and it is probable that in a state
of nature these leave the nest as quickly as possible. _Emenadia
flabellata_, a genus allied to _Metoecus_, has been discovered by Chobaut
to have a similar life-history, except that it attacks a solitary wasp of
the genus _Odynerus_.[139] An old record to the effect that a second
species of _Emenadia_, _E. bimaculata_, lives in the stalks of _Eryngium
campestre_, on the pith, is now thought to be erroneous. Fabre has found
the larvae and pupae of another Rhipiphorid in the cells of a bee,
_Halictus sexcinctus_.
The most remarkable of the Rhipiphorids, from the point of view of its
habits, is certainly _Symbius blattarum_, which is now treated as the same
as an Insect previously described by Thunberg from specimens found in amber
and called _Ripidius pectinicornis_. This species is parasitic in
cockroaches; the male and female are very different, the former being an
active winged Insect, while the female is worm-like, differing but little
from the larva, and never leaving the body of the cockroach. It is to be
regretted that the life-history is not better known. The species has been
found on board ship in vessels coming from India; the male has been met
with in several European countries, but the female is excessively rare.
FAM. 75. CANTHARIDAE OR MELOIDAE (_Blister-beetles_, _Oil-beetles_).-_-Head
with an abrupt neck; elytra and sides of the abdomen without any
coadaptation; each claw of the feet with a long appendage closely applied
beneath it._ This distinct family consists of Heteromera with soft
integument, and is remarkable for the fact that many of its members contain
a substance that when extracted {270}and applied to the human skin,
possesses the power of raising blisters. The life-history is highly
remarkable, the most complex forms of hyper-metamorphosis being exhibited.
The species now known amount to about 1500; there can be no difficulty in
recognising a member of the family by the above characters, except that in
a very few cases each claw bears a projecting tooth, instead of an elongate
appendage parallel with itself. The penultimate tarsal joint scarcely ever
broader than the preceding; the colour and style of markings are extremely
varied. There are two very distinct sub-families, Cantharides and Meloides;
the former are winged Insects, and are frequently found on flowers or
foliage. The Meloides are wingless, and consequently terrestrial; they have
a very short metasternum, so that the middle coxae touch the hind; and they
also have very peculiar wing-cases, one of the two overlapping the other at
the base; in a few Meloids the wing-cases are merely rudiments.
The post-embryonic development of these Insects is amongst the most
remarkable of modern entomological discoveries. The first steps were made
by Newport in 1851,[140] and the subject has since been greatly advanced by
Fabre, Riley, and others. As an example of these peculiar histories, we may
cite Riley's account[141] of _Epicauta vittata_ (Fig. 140), a
blister-beetle living at the expense of North American locusts of the genus
_Caloptenus_. The locust lays its eggs underground, in masses surrounded by
an irregular capsule, and the _Epicauta_ deposits its eggs in spots
frequented by the locust, but not in special proximity to the eggs thereof.
In a few days the eggs of the blister-beetle hatch, giving rise to little
larvae of the kind called triungulin (Fig. 140, A), because each leg is
terminated by three tarsal spines or claws. In warm, sunny weather these
triungulins become very active; they run about on the surface of the ground
exploring all its cracks, penetrating various spots and burrowing, till an
egg-pod of the locust is met with; into this the triungulin at once eats
its way, and commences to devour an egg. Should two or more triungulins
enter the same egg-pod, battles occur till only one is left.
{271}[Illustration: FIG. 140.—Hypermetamorphosis of _Epicauta vittata_.
North America. (After Riley.) A, Young larva or triungulin; B, Caraboid
instar or second larva; C, coarctate larva, or instar between the
Scarabaeoid and Scolytoid larva; D, Scarabaeoid larva, from which the
Scolytoid, or sixth, instar differs but little; E, pupa; F, imago.]
After a few days passed in devouring a couple of eggs, the triungulin sheds
its skin and appears as a different larva (Fig. 140, B), with soft skin,
short legs, small eyes, and different form and proportions; a second moult
takes place after about a week, but is not accompanied by any very great
change of form, though the larva is now curved, less active, and in form
like a larva of Scarabaeidae; when another moult occurs the fourth instar
appears as a still more helpless form of larva (Fig. 140, D), which
increases rapidly in size, and when full grown leaves the remains of the
egg-pod it has been living on, and forms a small cavity near by; here it
lies on one side motionless, but gradually contracting, till the skin
separates and is pushed down to the end of the body, disclosing a
completely helpless creature that has been variously called a semi-pupa,
pseudo-pupa, or coarctate larva (Fig. 140, C); in this state the winter is
passed. In spring the skin of the coarctate larva bursts, and there crawls
out of it a sixth instar which resembles the fourth (Fig. 140, D), except
in the somewhat reduced size and greater whiteness. It is worthy of remark
that the skin it has deserted retains its original form almost intact. In
this sixth instar the larva is rather active and burrows about, {272}but
does not take food, and in the course of a few days again moults and
discloses the true pupa (Fig. 140, E). As usual in Coleoptera this instar
lasts but a short time, and in five or six days the perfect beetle appears
(Fig. 140, F). It is extremely difficult to frame any explanation of this
complex development; there are, it will be noticed, no less than five
stages interposed between the first larval instar and the pupal instar, and
the creature assumes in the penultimate one a quasi-pupal state, to again
quit it for a return to a previous state. It is possible to look on the
triungulin and the pupal instars as special adaptations to external
conditions; but it is not possible to account for the intermediate instars
in this way, and we must look on them as necessitated by the physiological
processes going on internally. Nothing, however, is known as to these. It
may be well to mention that, after describing and figuring (_loc. cit._)
this series of instars, Riley changed his views as to their
nomenclature.[142] The following summary of the metamorphosis, to which we
have added the two nomenclatures of Riley—the original one, when different
from the amended one, being given in square brackets—may therefore be
useful, viz.—Egg; 1, triungulin-larva—moult; 2, Caraboid larva [second
larva, Caraboid stage]—moult; 3, Scarabaeoid larva [second larva,
Scarabaeoid stage]—moult; 4, Scarabaeoid larva [second larva, ultimate
stage] (large amount of food and much growth)—moult; 5, coarctate larva
[pseudo-pupa, or semipupa]; 6, Scolytoid larva [third larva] (active, but
little or no food taken)—moult; 7, pupa—moult; 8, perfect Insect.
M. Fabre has succeeded in elucidating the history of _Sitaris humeralis_, a
Cantharid that lives at the expense of bees of the genus _Anthophora_.[143]
The eggs of the _Sitaris_ are deposited in the earth in close proximity to
the entrances to the bees' nests, about August. They are very numerous, a
single female producing, it is believed, upwards of 2000 eggs. In about a
month—towards the end of September—they hatch, producing a tiny triungulin
of black colour; the larvae do not, however, move away, but, without taking
any food, hibernate in a heap, remaining in this state till the following
April or May, when they become active. Although they are close to the
abodes of the bees they do not enter them, but seek to attach themselves
{273}to any hairy object that may come near them, and thus a certain number
of them get on to the bodies of the _Anthophora_ and are carried to its
nest. They attach themselves with equal readiness to any other hairy
Insect, and it is probable that very large numbers perish in consequence of
attaching themselves to the wrong Insects. The bee in question is a species
that nests in the ground and forms cells, in each of which it places honey
and lays an egg, finally closing the receptacle. It is worthy of remark
that in the case of the _Anthophora_ observed by M. Fabre, the male appears
about a month before the female, and it is probable that the vast majority
of the predatory larvae attach themselves to the male, but afterwards seize
a favourable opportunity, transfer themselves to the female, and so get
carried to the cells of the bee. When she deposits an egg on the honey, the
triungulin glides from the body of the bee on to the egg, and remains
perched thereon as on a raft, floating on the honey, and is then shut in by
the bee closing the cell. This remarkable act of slipping on to the egg
cannot be actually witnessed, but the experiments and observations of the
French naturalist leave little room for doubt as to the matter really
happening in the way described. The egg of the bee forms the first
nutriment of the tiny triungulin, which spends about eight days in
consuming its contents; never quitting it, because contact with the
surrounding honey is death to the little creature, which is entirely
unfitted for living thereon. After this the triungulin undergoes a moult
and appears as a very different creature, being now a sort of vesicle with
the spiracles placed near the upper part; so that it is admirably fitted
for floating on the honey (Vol. V. Fig. 86, 10). In about forty days, that
is, towards the middle of July, the honey is consumed, and the vesicular
larva after a few days of repose changes to a pseudo-pupa (11 of the fig.
cited) within the larval skin. After remaining in this state for about a
month, some of the specimens go through the subsequent changes, and appear
as perfect Insects in August or September. The majority delay this
subsequent metamorphosis till the following spring, wintering as
pseudo-pupae and continuing the series of changes in June of the following
year; at that time the pseudo-pupa returns to a larval form (12 of the fig.
cited), differing comparatively little from the second instar. The skin,
though detached, is again not shed, so that this ultimate larva is enclosed
in two {274}dead skins; in this curious envelope it turns round, and in a
couple of days, having thus reversed its position, becomes lethargic and
changes to the true pupa, and in about a month subsequent to this appears
as a perfect Insect, at about the same time of the year as it would have
done had only one year, instead of two, been occupied by its metamorphosis.
M. Fabre employs the term, third larva, for the instar designated by Riley
Scolytoid larva, but this is clearly an inconvenient mode of naming the
instar. _Sitaris humeralis_ is now very rare in Britain, but it seems
formerly to have been more common, and it is not improbable that its
triungulin may have been the "_Pediculus melittae_," that was believed by
Kirby to be a sort of bee-louse. Some species of the genus _Meloe_ are
still common in Britain, and the Insects may be seen with heavy distended
abdomen grazing on herbage in the spring. The females are enormously
prolific, a single one producing, it is believed, about 10,000 eggs.
_Meloe_ is also dependent on _Anthophora_, and its life-history seems on
the whole to be similar to that of _Sitaris_; the eggs are, however, not
necessarily deposited in the neighbourhood of the bees' nests, and the
triungulins distribute themselves on all sorts of unsuitable Insects, so
that it is possible that not more than one in a thousand succeeds in
getting access to the _Anthophora_ nest. It would be supposed that it would
be a much better course for these bee-frequenting triungulins to act like
those of _Epicauta_, and hunt for the prey they are to live on; but it must
be remembered that they cannot live on honey; the one tiny egg is their
object, and this apparently can only be reached by the method indicated by
Fabre. The history of these Insects certainly forms a most remarkably
instructive chapter in the department of animal instinct, and it is a
matter for surprise that it should not yet have attracted the attention of
comparative psychologists. The series of actions, to be performed once and
once only in a lifetime by an uninstructed, inexperienced atom, is such
that we should _a priori_ have denounced it as an impossible means of
existence, were it not shown that it is constantly successful. It is no
wonder that the female _Meloe_ produces 5000 times more eggs than are
necessary to continue the species without diminution in the number of its
individuals, for the first and most important act in the complex series of
this life-history is accomplished by an extremely indiscriminating
instinct; the {275}newly hatched _Meloe_ has to get on to the body of the
female of one species of bee; but it has no discrimination whatever of the
kind of object it requires, and as a matter of fact, passes with surprising
rapidity on to any hairy object that touches it; hence an enormous majority
of the young are wasted by getting on to all sorts of other Insects; these
larvae have been found in numbers on hairy Coleoptera as well as on flies
and bees of wrong kinds; the writer has ascertained by experiment that a
camel's-hair brush is as eagerly seized, and passed on to, by the young
_Meloe_ as a living Insect is.
The histories of several other Cantharids have been more or less completely
discovered. Fabre has found the larva of _Cerocoma schaefferi_ attacking
the stores of provisions laid up by a fossorial wasp of the genus
_Tachytes_, and consisting of Orthoptera of the family Mantidae. The
student who wishes for further information may refer to M. Beauregard's
work on this family.[144]
Some half-dozen species of the genus _Cephaloon_ found in Siberia, Japan,
and North America, have, by some authorities, been separated as the family
Cephaloidae. Nothing is known as to the metamorphosis of these rare
beetles; and at present it is not necessary to distinguish them from
Cantharidæ.
FAM. 76. TRICTENOTOMIDAE.—_Large Heteromera, with powerful free projecting
mandibles; the antennae long, but with the terminal three joints short,
with angular projections on one side._ This family includes only two genera
and seven or eight species. They are very remarkable Insects; _Autocrates
aenea_ being three inches long. The family is of considerable interest, as
it seems to have no affinity with any other Coleoptera. The appearance of
the species somewhat reminds one of Lucanidae, or Prionides; but
Trictenotomidae have even less relation to those beetles than they have to
the members of the Heteromerous series. The Trictenotomidae appear to be
found only in the primitive forests of the Indian and Indo-Malayan regions.
Nothing is known as to their life-histories.
{276}SERIES V. PHYTOPHAGA.
_Tarsi apparently four-jointed, the three basal joints usually densely
set with cushion-like pubescence beneath; the third joint different in
form, being divided into two lobes, or grooved on its upper surface so as
to allow of the fourth joint being inserted near its base instead of at
its extremity. Head not forming a definite prolonged beak; its labrum
visible, the palpi rarely (and even then not completely) occluded in the
mouth._