Guide to the study of insects, and a treatise on those injurious and beneficial to crops: for the use of colleges, farm-schools, and agriculturists online

ward." He suggests that the development into a queen was
caused by the increased temperature of the queen-cell, above
that of the worker-cells.

But Messrs. F. Smith and Woodbury (Proceedings of the
Entomological Society of London, January 2, 1862) support F.



APIARI.E. 119

Huber's theory, that the change is due to "the quality as well
as quantity of food with which the royal larva is supplied,'*
though Dr. Leitch objects, that it has been by no means con-
clusively proved " that the so-called royal jelly differs in anv
respect from the ordinary food supplied to the worker larva;"
and Mr. AVoodbury cites the experiments of Dzierzon, as
quoted by Kleine, - that as Huber, by introducing some royal
jellv in cells containing worker-brood, obtained queens, it may
be possible to induce bees, to construct royal cells, when the
Apiarian prefers to have them, by inserting a small portion of
royal jelly in cells containing worker-larvae." Kleine takes " an
unsealed royal cell which usually contains an excess of
royal jelly and removes from it a portion of the jelly, on
the point of a knife or pen, and by placing it on the inner
margin of any worker cell, feels confident that the larvae in
them will be reared as queens."

Before these points are settled we must study the habits of
the Wild Bees, and of the other social Hymenoptera and White
Ants, together with the social Aphides more carefully. Mr. F.
\\ . Putnam pertinently states, "at present I cannot believe
that the peculiarity of food, or the structure of the cells, pro-
duces a difference of development in Humble-bees, for the lar-
vae, as has been previously stated, were seen to make their own
cells from the pollen paste. Is it not more natural to believe,
as lias been suggested to me by Professor J. Wyman, that the
dillereuce in the development of the eggs is owing to their be-
ing laid at various times after impregnation? Thus, if I am
right in supposing that the queens are impregnated by the
males late in the summer, the eggs, laid soon after, produce
the large queen larvae ; * the next set of eggs, laid in the spring,
produce the workers, or undeveloped females, while from those
deposited still later, male bees are principally developed."
(Proceedings of the Essex Institute, Salem, vol. iv, 1864, p.
103.)

Referring to Mr. Putnam's statement that there are both small
and large queens (besides the workers), Dr. Gerstaecker infers,

* Dr. Gerstiecker, on the other hand, states that " from the brood-cells of a nest
of Bombus musconnn, found by him on the 18th of September, there were devel-
oped at the end of the same month only workers."



120 HYMENOPTERA.

"from the examination of numerous individuals found flying in
the spring after hibernation, that these could not be considered
as true queens, since their ovaries were only moderately devel-
oped, though larger than those of the workers, while in the
true queen, captured in the summer, the ovaries were perfectly
developed. This corresponds almost entirely to what we find
in the wasps, whose spring females have only moderately de-
veloped ovaries."

How the Honey-bee builds its cells, and whether the} 7 are ex-
actly hexagonal, are questions that have interested the best
observers from Maraldi who wrote in 1712, and Reaumur,
whose Memoires appeared in 1740, down to the present date.
Their solution involves not only the closest observation of the
insect while at work, but also the shrewdest judgment to ex-
plain the facts observed and deduce a legitimate theory. Does
the bee intelligently plan her work out beforehand, or does she
follow the guidance of what is called instinct? Does she
construct hexagonal cells which are mathematically exact,
or does she vary the proportions of each cell, so that it is per-
fect only in its general ideal form ? Again, in making the cell,
is the bee actually capable of making such a cell alone, or is it
due to the resultant action of several bees? Professor J. Wy-
man is of the latter opinion, as he thinks "that if left alone to
build a single cell, this would most probably be round. In the
cells of Melipona, as Huber's plate shows, they are only hex-
agonal when in contact with the adjoining cells." (Proceed-
ings of the Boston Society of Natural History, x, p. 278,
1866.)

A similar view is that proposed in 1862 by the Rev. Samuel
Haughton, in a paper read before the Natural History Society
of Dublin, where he says, according to Mr. F. Smith, that the
hexagonal form of the cell " may be accounted for simply by the
mechanical pressure of the insects against each other during
the formation of the cell. In consequence of the instinct that
compels them to work with reference to a plane, and of the
cylindrical form of the insect's body, the cells must be hex-
agonal."

Mr. G. R. Waterhouse (Transactions of the Entomological
Society of London. Third series, vol. ii, p. 129, 1864) has



APIARI^E. 121

proposed what has been called the ''circular theory," or what
the author himself terms the principle of working in seg-
ments of circles." He contends " that the hexagonal form of the
cells of certain bees and wasps may, and does, arise out of this
mo. ! of action when under certain conditions ; that those condi-
tions arc, that the cells are so commenced that their natural cir-
cumferences, as the work proceeds, are either simply brought
into contact with each other, or that the cells are so placed that
the (we will say theoretical) circumferences must intersect.
Contact with adjoining cells, then, is an essential condition to
bring about the hexagonal form as I have before pointed out
(See Proceedings of the Entomological Society, 1858, p. 17) ;
but for this result it is not necessary that a hexagonal cell
should be completely surrounded by other cells."

Is not this theory, after all, too mechanical ? Is not our bee
more of a free agent ? Does it not have a mind to design its
work? Mr. F. Smith, who has devoted years to the study of
Hymenoptera, especially the higher forms of this suborder, the
Bees and Wasps, replies to both theories of Waterhouse and
Haughton, by bringing in the case of the Wasps which also
build hexagonal cells, showing that a solitary wasp will build
its cells in very regular hexagons. Thus the nest of the soli-
tary Wasp, /can a guttatipennis, "consists of a double row, the
number of cells being ten ; I now direct your attention to the
fact that all the cells are perfectly hexagonal, the exterior
planes being as beautifully finished as those in contact with
the inner planes of the opposing cells. I have placed a draw-
ing of this nest (Plate 5, Fig. 7) in the box on the table, and I
particularly wish you to observe, that the first cell is carried
up in a perfectly hexagonal form above the adjoining cells ; a
proof that, if Wasps never build perfect isolated hexagonal cells,
they certainly possess the capability of doing so. The exterior
of all the cells, as I before observed, is hexagonal, not cylindri-
cal, until fresh cells are added on the outer side, as was ob-
served to be the case in combs of the Hive-bee, by Mr.
Tegetmeier." (Proceedings of the Entomological Society of
London. Third series, ii, 1864, p. 135.)

An examination of the cells of three species of Polistes (the
female of which begins alone in the spring to build her nest,



122 HYMENOPTERA.

the cells of which are afterwards greatly increased in number
after the first brood of females appear) , convinced us that
the Wasp begins with the circular cup -shaped form of cell, and
when about depositing an egg in it, changes her mode of ope-
rating, builds up the edges into a hexagonal form, and carries
up the rim of each cell independently to its required height.
She thus apparently changes her plan at a certain stage of the
work, and is so far a free agent.

Mr. Smith also exhibited a portion of the nest of another
wasp, Tatua Mono (Plate 5, Fig. 9), that proved to his mind
the primary intention of the wasp instinctively to build cells
with exactly six sides. The figure represents part of one of
the flat floors, on which the foundations of the cells are laid in
regular hexagons, instead of beginning in hemispherical cups.

Mr. Smith (p. 141) concludes, "that all hexagonal cells are
not constructed upon a circular principle, and that the primary
idea of all social bees and wasps is not to produce cylindrical
cells with hemispherical bases."

In this connection the following extract from Mr. Smith's
remarks is of interest : "It may not be known that in order to
expedite the building of honey-combs, it is a common practice
with bee-keepers in Germany to furnish hives with artificial
foundations for the cells ; these consist of sheets of wax, upon
which is impressed a series of pyramidal hollows ; in fact, the
counterpart of a comb built by the bees themselves, entirely
deprived of the cell-walls ; and it is from such a piece of comb
that the casts for the artificial foundations are obtained. A
piece of casting of this description I lay before you, and I par-
ticularly call your attention (addressing the members of the
Entomological Society of London) to the commencement of the
outer cells ; you will see, in some instances, a single plane of
the hexagonal cell commenced, in others two or. three are in
progress ; here you have a ground-plan supplied, or, I may say,
the foundations of the habitations ready prepared, upon which
the laborers are to raise the walls, and you may see how admi-
rably they have done it. Instinct enables the bee to construct
hexagonal cells without teaching, and, we are told, in one un-
deviating manner. Surely the example before us exhibits an
amount of intelligence on the part of the bees in availing them-



12.;

selves of such adventitious aid. Must we not henceforth,
when spi-:ikin<r of the marvels of the hive or the vespiary, erase
from our vocabulary such terms as blind instinct ; and must we
not cease to stigmatize the bee as a mere machine?"

At the meeting of the same society held Feb. 1, 1864, Mr.
F. Smith exhibited a collection of Wasps' nests, one of Vespa
r nf< i, the rest of V. vulgaris; they were in various stages of
formation, the earliest consisting of only a single cup contain-
ing the first egg, others consisting of three or four cups, whilst
others again were more complete. The whole had been arti-
ficially obtained by Mr. Stone, who tempted the wasps to build
by excavating holes in banks and furnishing them with foot-
stalks ; in fact, Mr. Stone appeared to possess the power of
inducing wasps to build nests of almost any shape he
pleased.

But to return to the cell of the Bee. It should first be
proved that the cells are not exactly and mathematically per-
fect hexagons, though sufficiently so for the purpose for which
they are used. In the Proceedings of the American Academy
of Arts and Sciences, vol. vii, 18G6, Professor "VVyman has, by
a most careful as well as novel and ingenious mode of investiga-
tion, proved that the cells are all more or less imperfect, and
that a hexagonal cell mathematically exact, does not exist in
nature, but only in theory.

The form of the cell is liable to marked variations, chief
among which the following may be mentioned, in the author's
own words :

"1. The diameters of workers' cells may so vary, that ten
of them may have an aggregate deviation from the normal
quantity equal to the diameter of a cell. The average varia-
tion is a little less than one half that amount, namely, nearly
0.10 inch, in the same number of cells.

"2. The width of the sides varies; and this generally in-
volves a variation of the angles which adjoining sides make
with each other, since the sides vary not only in length but in
direction.

"3. The variation in the diameters does not depend upon
accidental distortion, but upon the manner in which the cell
was built.



124



IIYMENOPTERA.



"4. The relative size of the rhombic faces of the pyramidal
base is liable to frequent variation, and this where the cells are
not transitional from one kind to another.

U 5. When a fourth side exists in the basal pyramid, it may
be in consequence of irregularity in the size of the cells, or of
incorrect alignment of them on the two sides of the comb."

Sometimes one of the faces is lost, and a new one formed,
so that all the basal portion of the cell becomes reversed, as
ABC w ill be seen by refer-
ence to Figs. 73 and
74 ; the first repre-
Fig. 73. senting the cells when

the base is viewed, and the second when looked at perpendic-
ularly to one of the sides. In both figures A indicates the
ordinary form of the cell. The whole ABC
series of Fig. 74 shows the gradual
introduction of the new face, which
is seen on the lower border, and the
elimination of one of the original faces,
which is seen on the upper border. At
B, which is intermediate between the
two extremes, the four faces consist of two equal rhombs,
one of which is the outgoing and the other the incoming one,
A and two equal hexagons. B, Fig.

74, represents the sides of the same
cell, which, instead of forming three
trapeziums, as at A, a, 6, c, now
form two pentagons, a! and c', and a
parallelogram, b'. At C, Figs. 73
and 74, the forms are in all respects
the reverse of those of A. A and C
are symmetrical with each other, and
B is symmetrical in itself. No pre-
cise number of cells is necessary
for the purpose of making this transition, for it may take
place in two or three, or extend* through a long series, as in
Fig. 73.

"6. Ordinarily, the error of alignment does not amount to
more than one or two diameters of a cell. But occasionally



OS V

Fig.




C

a



Fig. 75.



APIARLE. 125

the rows of cells on one side of the comb may deviate from

their true direction with regard to those on the other, to the

extent of 30."

u Thus, if a piece of normal comb be held in the position in

which it was built, two of the opposite angles of the hexagon,

Fig. 75, A, a, will be in the

same vertical line, and two

of the sides will be parallel

to this. The same is true

of the opposite side of the

comb ; and thus all the cor- * "11 111 r~/

responding parts of the cells

on the two sides will be par-
allel. In the deviation we

are now noticing, the change

is like that represented in A,

where the cell a is in its

true position, while the cell

6, which is from the oppo-
site side, 'and is in contact

with a, varies from it by

about 30. If we look at

the'se two cells in the direc-
tion of their sides as at B,

the prism a will have one Fig. 76.

of its angles towards the eye, and b one of its sides.

In consequence of this deviation and the continual crossing

of the rows on opposite sides, the pyramidal base is not made,

and the cell is shortened.

"7. In curved or bent combs the
cells on the concave side tend to be-
come narrower, while those on the
other tend to become broader to-
wards their mouths. In Fig. 7G
(this and Figs. 77 and 78 are made
Fi - 77> from impressions obtained directly

from the comb and transferred to wood ; they represent the

form of the cells exactly), as in the central line of cells, there

are a variety of hexagons, each resulting from the union





126



HYMENOPTERA.



of two cells, the base being double while the mouth is
single. That on the line a, 6, has three sides at one end,
united by two long sides with one at the other, and thus two
of the opposite sides are not parallel ; at c, cZ, two sides at
a b




'9



c d e f

Fig. 78.

either end are united by two long sides, these last being par-
allel ; and at e, /, the mouth of the compound cell has seven
sides. Each has a partition at its base, separating the two
originally distinct cells, and each was lined w r ith a cocoon,
showing that it had been used for rearing young. At #, not
only has the partition between the combining cells disappeared,
but also three of the sides of each cell."

The bees do not appear to have any systematic way of mak-
ing a transition from worker to drone cells, which are one-fifth
larger than the former. More commonly, they effect it by a
gradual alteration of the diameters, thus enlarging a worker
into a drone, or narrowing a drone into a worker cell. This
alteration is usually made in from four to six rows. In one case



APIAHI^E. 127

Professor "Wyman noticed the transition made with only one
cell, as in Fig. 78, but not without destroying the regularity of
the two adjoining rows.

u ln consequence of the gradual narrowing or widening of
the transition cells, the comb tends to become more or less tri-
angular and the cells to become disturbed. The bees counter-
act this tendency by the occasional intercalation of an additional
row, of which two instances are given in Fig. 78, at a and &,
where three rows of worker cells are continuous with two of
drone cells, c, d and e, /; or, reversing the statement, and
supposing the transition, as in the building of the comb, is
from worker to drone-cells, a row of the latter is from time to
time omitted as the rows a and b; in this way, the regularity of
the comb is preserved."

Honey-cells are formed either by enlarging the ordinary
brood-cells, or adding them to others often larger, or by con-
structing a new comb, devoted entirely to the storing of honey.
"While the cells of this last are built unequivocally in accord-
ance with the hexagonal type, they exhibit a range of variation
from it which almost defies description."

No Ichneumon-flies are known to attack the larva of the
Honey-bee, nor in fact, with few exceptions, any of the wild
bees, owing, probably, to the difficulty of their gaining access
to them, since Anomalon vesparum has been reared from the
cells of wasps which are more exposed than those of bees.
But the Honey, as well as the wild bees, are afflicted by a
peculiar assemblage of insect-parasites, some of which have
the most remarkable habits. The most formidable pest of the
Hive-bee is the Bee Fly, Phora incrassata, which in Europe
sometimes produces the well-known disease called " foul-
brood." The Bee-louse, Braula cceca, is, in Europe, sometimes
troublesome to the adult bee, while Trichodes apiarius, a beetle,
devours the larva*. The larvae of Meloe and Stylops are known
in Europe to infest the Honey-bee, and among the low intesti-
nal worms Assmusi enumerates Gordius subbifurcus which in-
fests the drones of the Honey-bee as well as other insects.
Professor Siebold has also described Mermis albicans, which
is a similar kind of hair-worm, from two to five inches long,
and whitish in color. This worm is also found, strangely



128 HYMENOPTERA.

enough, only in the drones, though it is the workers which
frequent watery places (where the worm deposits its eggs) to
appease their thirst. The Wax-moths, Galleria cereana and
Achroia alvearia, do much harm by consuming the wax and
thus breaking down the cells, and by filling the hive with
their webs.*

The genus Apis is indigenous in "South America, though the
Honey-bee has been extensively introduced into the West In-
dies. Our Honey-bee is replaced in the tropics by the stingless,
minute bees, which store up honey and live in far more numer-
ous colonies. The cells of Melipona are hexagonal, nearly
approaching in regularity those of the Hive-bee, while the
honey-cells are irregular, much larger cavities, which hold about
one-half as much honey as a cell of the Humble-bee. From a
paper on the Brazilian Honey-bees, read by Mr. F. Smith be-
fore the Entomological Society of London, March, 1863, he
states that the Meliponas are small insects, having wings shorter
than the abdomen, the latter being very convex and oblong ;
their mandibles never being dentate ; while the Trigonas have
the wings more ample, and longer than the abdomen, which is
short, somewhat triangular, while the mandibles are serrated,
denticulate, or sometimes edentate. The Meliponas are re-
stricted to the new world, while Trigona extends into Africa,
India, and Australasia.

"All these bees are honey gatherers, but the honey collected
by the different species varies greatly in quality : from the
nests of some it is excellent ; from others, worthless. The
honey of the species i Mombuca' is said to be black and sour,
the quality being dependent on species of flowers from which
the honey is collected. This great difference in the honey of
the various species is apparently confirmatory of the fact that
each species confines itself to particular flowers, never visiting
any other kind. The different relative length of the tongue in

* EXPLANATION OP PLATE 2. Parasites of the Honey-bee. Fig. 1, Pliora incras-
sata; Fig. 2, pupa; Fig. 3, larva. Fig. 4, Braula cceca; Fig. 5, larva. Fig. 6, Tri-
chodes apiarius . a, larva ; 6, pupa. Fig. 7, Meloe angusticollis ; Fig. 8, fveshly hatched
larva; Fig. 9, second stage of larva; Fig. 10, first stage of semi-pupa; Fig. 11,
pupa. Fig. 12, Stylops Childreni in the body of a wild bee, Andrena; Fig. 13, top
view of the same removed from its host; Fig. 14, male of the same; a, side view.
Fig. 15, Mucor mellitophorus, a parasitic fungus. Fig. 16, unknown larva found in
nest of Humble-bee. Descriptions of the insect parasites will be given beyond.



APIARI^E. 129

the species is also confirmatory of the same supposition ; in-
deed, the great diversity in this respect observable in these
bees, appears to me to be analogous to a similar diversity in the
length of the bills of humming-birds, which, it is well known,
are always adapted for reaching the nectaries of the particular
flowers which they usually frequent."

In regard to the immense numbers of individuals in a col-
ony, Mr. Stretch, who collected them at Panama, "found a
nest several feet in length in the hollow of a tree, containing
thousands of individuals, their numbers being, as he informs
me, apparently countless.

"Gardner, in his travels, gives a list of such species (of
Melipona) as he met in the provinces of Piauhy and Goyaz,
where he found them numerous ; in every house, he says, 'you
find the honey of these bees ; ' many species, he tells us, build in
the hollow trunks of trees, others in banks ; some suspend
their nests from branches of trees, whilst one species constructs
its nest of clay, it being of large size ; the honey of this spe-
cies, he says, is very good." (Smith.)

In a nest of Trigona carbonaria from Eastern Australia,
Smith, of the British Museum, found from 400 to 500 dead
workers crammed in the spaces between the combs, but lie
did not find a female among them. The combs are arranged
precisely similar to those of the common wasp. The number of
honey-pots, which are placed at the foot of the nest, amounted
to 250.

Smith inclines to the opinion that the hive of Trigona con-
tains several prolific females ; "the accounts given of the mul-
titudes inhabiting some nests is too great, I think, to render it
possible that one female could produce them all. Mr. Stretch
described a hive that he saw, occupying the interior of a decay-
imr tree, that measured six feet in length, and the multitude of
bees he compared to a black cloud. M. Guerin found six fe-
males in a nest of Melipona fulvij)es"

Hill states, in Gosse's Naturalist's Sojourn in Jamaica,
"that the wax of these bees [Trigona] is very unctuous and
dark colored, but susceptible of being whitened by bleaching.
The honey is stored in clusters of cups, about the size of
pigeon's eggs, at the bottom of the hive, and always from the
9



130 HYMENOPTEEA .

brood-cells. The brood-cells are hexagonal ; they are not
deep, and the young ones, when ready to burst their casement, '
just fill the whole cavity. The mother bee is lighter in color
than the other bees, and elongated at the abdomen to double
their length." Smith also states that the female of this genus
has the abdomen greatly distended, reminding one of the
gravid female of the White Ant. (Smith, Proc. Ent. So'c.,
London, Dec. 7, 18G3.)

In North America, our nearest ally, as regards its habits, of
the true Honey-bee, is the Humble-bee (Bombus), of which
over forty species are known to inhabit North America.

The economy of the Humble-bee is thus : the queen awakens
in early spring from her winter's sleep beneath the leaves or
moss, or in deserted nests, and selects a nesting-place generally
in an abandoned nest of a field-mouse, or beneath a stump or
sod, and "immediately," according to Mr. F. W. Putnam,
"collects a small amount of pollen mixed with honey, and in
this deposits from seven to fourteen eggs, gradually adding to
the pollen mass until the first brood is hatched. She does not