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The larvae (for figure of larva of Bombus, see Vol. V. p. 488) are,
without known exception, legless grubs, of soft consistence, living
entirely under cover, being protected either in cells, or, in the case of
social Hymenoptera, in the abodes of the parents. The larvae of Ants
and fossorial Hymenoptera have the anterior parts of the body long
and narrow and abruptly flexed, so that their heads hang down in a
helpless manner. All the larvae of Aculeates, so far as known, are
remarkable from the fact that the posterior part of the alimentary
canal does not connect with the stomach till the larval instar is more
or less advanced; hence the food amongst which they live cannot be
sullied by faecal matter. The pupa is invariably soft, and assumes
gradually the colour of the perfect Insect. Almost nothing is known as
to the intimate details of the metamorphosis, and very little as to the
changes of external form. According to Packard a period intervenes
between the stadium of the full-grown larva and that of the pupa, in
which a series of changes he speaks of as semi-pupal are passed
through; these, however, have not been followed out in the case of
any individual, and it is not possible to form any final idea about
them, but it seems probable that they are largely changes of external
shape, in conformity with the great changes going on in the internal
organs. Owing to the fragmentary nature of observations, much
obscurity and difference of opinion have existed as to the
metamorphosis of Aculeate Hymenoptera. Sir S. Saunders gives the
following statement as to the larva of a wasp of the genus
Psiliglossa,[5] just before it assumes the pupal form: "The respective
segments, which are very distinctly indicated, may be defined as
follows:—The five anterior, including the head, are compactly welded
together, and incapable of separate action in the pseudo-pupa state;
the third, fourth, and fifth bearing a spiracle on either side. The
thoracical region terminating here, the two anterior segments are
assignable to the development of the imago head, as pointed out by
Ratzeburg." This inference is not, however, correct. We have seen
that in the perfect Insect of Petiolate Hymenoptera the first
abdominal segment is fixed to the thorax, and Saunders' statement
is interesting as showing that this assignment of parts already exists
in the larva, but it in no way proves that the head of the imago is
formed from the thorax of the larva. It has been stated that the larvae
of the Aculeata have a different number of segments according to
the sex, but this also is incorrect. The difference that exists in the
perfect Insects in this respect is due to the withdrawal of the terminal
three segments to the interior in the female, and of two only in the
male. The larva consists of fourteen segments, and we find this
number distributed in the female perfect Insect as follows: one
constitutes the head, four segments the thorax and propodeum,
followed by six external segments of the restricted abdomen, and
three for the internal structures of the abdomen. This agrees with
Forel's statement that in the ants the sting is placed in a chamber
formed by three segments.
The development of the sting of the common bee has been studied
by Dewitz.[6] It takes place in the last larval stage. Although nothing
of the organ is visible externally in the adult larva, yet if such a larva
be placed in spirit, there can be seen within the skin certain small
appendages on the ventral surface of the penultimate and
antepenultimate abdominal segments (Fig. 4, A) placed two on the
one, four on the other; these are the rudiments of the sting. In the
course of development the terminal three segments are taken into
the body, and the external pair of the appendages of the twelfth body
segment (the ninth abdominal) become the sheaths of the sting, and
the middle pair become the director; the pair of appendages on the
eleventh segment give rise to the needles or spiculae. The sting-
rudiments at an earlier stage (Fig. 4, C) are masses of hypodermis
connected with tracheae; there is then but one pair on the twelfth
segment, and this pair coalesce to form a single mass; the rudiments
of the pair that form the director are differentiated secondarily from
the primary pair of these masses of hypodermis. A good deal of
discussion has taken place as to whether the component parts of the
sting—gonapophyses—are to be considered as modifications of
abdominal extremities (i.e. abdominal legs such as exist in
Myriapods). Heymons is of opinion that this is not the case, but that
the leg-rudiments and gonapophysal rudiments are quite distinct.[7]
The origin of the sting of Hymenoptera (and of the ovipositor of
parasitic Hymenoptera) is very similar to that of the ovipositor of
Locusta (Vol. V. p. 315 of this work), but there is much difference in
the history of the development of the rudiments.
Fig. 4—Development of sting of the bee: A and C, ventral; B, side view.
A, End of abdomen of adult larva: a, b, c, d, the last four
segments, c being the eleventh body segment, 11; b bearing two
pairs, and c one pair, of rudiments. B, Tip of abdomen of adult
bee: 9, the ninth, d, the tenth body segment. C, Rudiments in the
early condition as seen within the body: c, first pair; b, the second
pair not yet divided into two pairs; b″, c′, commencement of
external growths from the internal projections. (After Dewitz.)
We shall here follow the usual method of treating all the fossorial
wasps as forming a single group, uniting Ashmead's Entomophila
and Fossores, as we think their separation is only valid for the
purposes of a table; the Pompilidae placed by the American savant
in Fossores being as much allied to Entomophila as they are to the
other Fossores with which Ashmead associates them.
There are about 150 genera and 1500 species of bees at present
known. Some call the division Mellifera instead of Anthophila. The
term Apidae is used by some authorities to denote all the bees, while
others limit this term to one of the families or sub-divisions. The bees
are, as a rule, distinguished from other Hymenoptera by the hairs, by
the great development of the mouth parts to form a proboscis
(usually, but not correctly, called tongue), and by the modification of
the hind-legs; but these distinctive characters are in some of the
species exhibited in so minor a degree of perfection that it is not
easy to recognise these primitive forms as Anthophila. A few general
remarks on the three points mentioned will enable the student to
better appreciate the importance of certain points we shall
subsequently deal with.
The bees are, as a rule, much more covered with hair than any other
of the Hymenoptera. Saunders[10] states that he has examined the
structure of the hairs in all the genera of British Aculeata, and that in
none but the Anthophila do branched and plumose hairs occur. The
function of this kind of hairs is unknown; Saunders suggests[10] that
they may be instrumental in the gathering of pollen, but they occur in
the parasitic bees as well as in the males, neither of which gather
pollen. The variety of the positions they occupy on the body seems
to offer but little support to the suggestion. Not all the hairs of the
bee's body are plumose, some are simple, as shown in Fig. 5, A,
and this is specially the case with the hairs that are placed at the
edges of the dilated plates for carrying pollen. In some forms there is
an extensive system of simple hairs all over the body, and the
"feathers" are distributed between these; and we do not see any
reason for assuming that the feathered are superior to the simple
hairs for gathering and carrying pollen. Some bees, e.g. Prosopis,
Ceratina, have very little hair on the body, but nevertheless some
plumose hairs are always present even though they be very short.
Fig. 6—A, Worker of the honey-bee (Apis mellifica), with pollen plates
laden; B, basal portions of a middle-leg (trochanter with part of
coxa and of femur) with plumose hairs and grains of pollen; C,
one hair bearing pollen-grains.
This description will suffice for present purposes, as the other parts
of the mouth will be readily recognised by the aid of figure 9, A, B, C.
In the exquisitely endowed South American genus Euglossa (Fig.
18), the proboscis is somewhat longer than the whole of the body, so
that its tip in repose projects behind the body like a sting.
The correct nomenclature of the parts connected with the lower lip is
not definitely settled, authorities not being agreed on several points.
The whole of the proboscis is usually called the tongue; this,
however, is admittedly an erroneous application of this term. The
terminal delicate, elongate, flexible organ is by some called the
tongue; but this again is wrong: the lingua in Insects is the
hypopharynx; this part is developed in a peculiar manner in bees, but
as it is not tongue-like in shape, the term lingua is not suitable for it,
and should be dismissed altogether from the nomenclature of the
bee's trophi; it is used at present in two different senses, both of
which are erroneous. We see no objection to describing the flexible
apical portion of the proboscis as the ligula. The lorum is probably a
special part peculiar to the higher bees; according to Saunders it is
not present as a specialised part in some of the primitive forms.[14]
The application of the terms mentum, submentum and hypoglottis is
open to the same doubts that exist with regard to them in so many
other Insects, and we have omitted the term hypoglottis altogether,
though some may think the mentum entitled to that name.
The way in which the proboscis of the bee acts has been very largely
discussed, with special reference to the question as to whether it is a
sucking or a licking action. It is impossible to consider either of these
terms as applicable. The foundation of the action is capillary
attraction, by which, and by slight movements of increase and
contraction of the capacity of various parts, the fluid travels to the
cavity in front of the hypopharynx: here the scales of the maxillae
leave a vacant space, (Fig. 7, e) so that a cup or cavity is formed,
the fluid in which is within reach of the tip of the dependent
epipharynx (c), which hangs down over the front of the hypopharynx
(and is so shaped that its tip covers the cup); it is between these two
parts that the fluid passes to reach the pharynx. It is no doubt to
slight movements of the membranous parts of the hypopharynx and
of the epipharynx that the further progress of the nectar is due, aided
by contraction and expansion of the pharynx, induced by muscles
attached to it. It should be recollected that in addition to the
movements of the head itself, the hypopharynx is constantly
changing its dimensions slightly by the impulses of the fluid of the
general body cavity; also that the head changes its position, and that
the proboscis is directed downwards as well as forwards. Those who
wish to pursue this subject should refer to the works of Breithaupt[15]
and Cheshire.
The other external characters of the Bees call for little remark. The
pronotum is never very large or much prolonged in front, and its hind
angles never repose on the tegulae as they do in the wasps,[16] but
extend backwards below the tegulae. The hind body is never
narrowed at the base into an elongate pedicel, as it so frequently is
in the Wasps and in the Fossors; and the propodeum (the posterior
part of the thorax) is more perpendicular and rarely so largely
developed as it is in the Fossors; this last character will as a rule
permit a bee to be recognised at a glance from the fossorial
Hymenoptera.
The buzzing of bees (and other Insects) has been for long a subject
of controversy: some having maintained that it is partially or wholly
due to the vibration of parts connected with the spiracles, while
others have found its cause in the vibrations of the wings. According
to the observations of Pérez and Bellesme,[18] two distinct sounds
are to be distinguished. One, a deep noise, is due to the vibration of
the wings, and is produced whenever a certain rapidity is attained;
the other is an acute sound, and is said to be produced by the
vibrations of the walls of the thorax, to which muscles are attached;
this sound is specially evident in Diptera and Hymenoptera, because
the integument is of the right consistence for vibration. Both of these
observers agree that the spiracles are not concerned in the matter.
The young of bees are invariably reared in cells. These (except in
the case of the parasitical bees) are constructed by the mothers, or
by the transformed females called workers. The solitary bees store
the cells with food, and close up each cell after having laid an egg in
it, so that in these cases each larva consumes a special store
previously provided for it. The social bees do not close the cells in
which the larvae are placed, and the workers act as foster-mothers,
feeding the young larvae after the same fashion as birds feed their
nestling young. The food is a mixture of honey and pollen, the mixing
being effected in various ways and proportions according to the
species; the honey seems to be particularly suitable to the digestive
organs of the young larvae, and those bees that make closed cells,
place on the outside of the mass of food a layer more thickly
saturated with honey, and this layer the young grub consumes
before attacking the drier parts of the provisions. The active life of
the larva is quite short, but after the larva is full-grown it usually
passes a more or less prolonged period in a state of quiescence
before assuming the pupal form. The pupa shows the limbs and
other parts of the perfect Insect in a very distinct manner, and the
development of the imago takes place gradually though quickly.
Some larvae spin cocoons, others do not.
The more important of the groups that have been recognised are—
(1) the Obtusilingues, short-tongued bees, with the tip of the lingua
bifid or broad; (2) Acutilingues, short-tongued bees, with acute tip to
the tongue; these two groups being frequently treated of as forming
the Andrenidae. Coming to the Apidae, or the bees with long and
folded tongues, there have been distinguished (3) Scopulipedes,
bees carrying pollen with their feet, and (4) Dasygastres, those that
carry it under the abdomen; some of the parasitic and other forms
have been separated as (5) Denudatae (or Cuculinae); the Bombi
and the more perfectly social bees forming another group, viz. (6)
Sociales. A group Andrenoides, or Panurgides, was also proposed
for certain bees considered to belong to the Apidae though exhibiting
many points of resemblance with the Andrenidae. This arrangement
is by no means satisfactory, but as the tropical bees have been but
little collected, and are only very imperfectly known, it is clear that we
cannot hope for a better classification till collections have been very
much increased and improved. The arrangement adopted in Dalla
Torre's recent valuable catalogue of bees[19] recognises no less than
fourteen primary divisions, but is far from satisfactory.
Fig. 10—Prosopis signata. Cambridge. A, Female; B, front of head of
female; C, of male.
The two genera Prosopis and Sphecodes have been recently formed
into a special family, Archiapidae, by Friese,[20] who, however,
admits that the association is not a natural one. The term should be
limited to Prosopis and the genera into which it has been, or shortly
will be, divided. The primitive nature of the members of this genus is
exhibited in all the external characters that are most distinctive of
bees; the proboscis (Fig. 9, B, C), is quite short, its ligula being very
short, and instead of being pointed having a concave front margin.
The body is almost bare, though there is some very short feathered
plumage. The hind legs are destitute of modifications for industrial
purposes. Owing to these peculiarities it was for long assumed that
the species of Prosopis must be parasites. This is, however, known
not to be the case so far as many of the species are concerned.
They form cells lined with a silken membrane in the stems of
brambles and other plants that are suitable, or in burrows in the
earth, or in the mortar of walls; individuals of the same species
varying much as to the nidus they select. The food they store in
these cells is much more liquid than usual, and has been supposed
to be entirely honey, since they have no apparatus for carrying
pollen. Mr. R. C. L. Perkins has, however, observed that they
swallow both pollen and nectar, brushing the first-named substance
to the mouth by aid of the front legs. He has ascertained that a few
of the very numerous Hawaiian species of the genus are really
parasitic on their congeners: these parasites are destitute of a
peculiar arrangement of hairs on the front legs of the female, the
possession of which, by some of the non-parasitic forms, enables
the bee to sweep the pollen towards its mouth. These observations
show that the structural peculiarities of Prosopis are correlative with
the habits of forming a peculiar lining to the cell, and of gathering
pollen by the mouth and conveying it by the alimentary canal instead
of by external parts of the body. Prosopis is a very widely distributed
genus, and very numerous in species. We have ten in Britain;
several of them occur in the grounds of our Museum at Cambridge.
The species of the genus Colletes are hairy bees of moderate size,
with a good development of hair on the middle and posterior femora
for carrying pollen. They have a short, bilobed ligula like that of
wasps, and therein differ from the Andrenae, which they much
resemble. With Prosopis they form the group Obtusilingues of some
taxonomists. They have a manner of nesting peculiar to themselves;
they dig cylindrical burrows in the earth, line them with a sort of
slime, that dries to a substance like gold-beater's skin, and then by
partitions arrange the burrow as six to ten separate cells, each of
which is filled with food that is more liquid than usual in bees. Except
in regard to the ligula and the nature of the cell-lining, Colletes has
but little resemblance to Prosopis; but the term Obtusilingues may be
applied to Colletes if Prosopis be separated as Archiapidae. We
have six species of Colletes in Britain.