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Psyche 7:19-26, 1894.

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Psyche, Vol. VU, 1894.
Plate I.
AFRICAN NOCTUE, (HOLLAND.)
(REDUCED ONE-FI~.)




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PSYCHE.
THE HABITS OF THE ACULEATE HYMENOPTERA.-I. BY WILLIAM H. ASHMEAD, WASHINGTON, D. C. [Annual address of the retiring president of the Cambridge Entomological Club, 12 January, 1894.1 The subject of my address is one that
has been rarely touched by American
entomologists, although offering one
of the most attractive and richest fields for research and discovery, as connected with it are many problems of biologic
.
and philosophical importance, which if
solved, would throw much light upon
many of the moted questions of the day
-evolution of species, development of
sexes, specialization of organs, trans-
mission of acquired characters, adapta-
bility to environment, etc.
The first American to publish any-
thing on the subject was John Bartram,
who published several articles : the first entitled "An account of some curious
wasp-nests made of clay," was published
as early as 1745 (Phil. trans., vol. 43, pp. 363-368) ; the second, "A clescrip-
tion of the great black wasp of Penn."
(1. c., vol. 46, 1750, PP. 278-280) ; the third, "On the Yellow wasp of Perm."
(1. c., vole 53, 1763, pp. 57-39).
This last paper is of the deepest in-
terest as it evidently refers to the habits of a Bembecid, and the accounts of
which, now after over a century and a
quarter, have only recently been con-
firmed, in Europe, by the observations
of Fabre and Wesenberg on a similar
fossorial wasp, Bedex rostrata Fabr.
From John Bartram to our next
writer, Benjamin Henry Latrobe, who
wrote a paper entitled L"l'wo species of the Sphex or wasp found in Virginia
and Penn." (Phil. soc., vol. 6, 1809, p. 73) is an unbroken period of nearly half a century. Then we have a period of
longer or shorter intervals, with con-
tributions from Thomas Say, Dr. T. W.
Harris, F. W. Putnam, Dr. Lincecum ,
Dr. A, S. Packard, Wm. Couper, Ben-
jamin D. Walsh, Prof. C. V. Riley,
E. Baynes Reed, L. 0. Howard,
Frederick V. Coville, Charles Robert.
son, C. L. Marlatt, and Dr. A.
Davidson.
It is now, I believe, almost univer-
sally conceded by all students, who
have given any study at all to the acule- ate Hymenoptera, that among them are
to be found the most specialized, highly developed and intelligent insects. In
fact, the marvellous intelligence exhib- ited by many of the species in this order, in their social habits, the structure of their nests, care of their young, etc., has from time immemorial attracted the
attention of man, and in both ancient
and modern literature many allusions
to them may be found.
It is surprising, therefore, that so
many centuries have past and so little




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20 P5'X'h723. [February 1894.
comparatively is known of the vast
majority of the most common forms.
It is hoped that a few new facts
respecting the habits of some of our
species will be found in this address,
but it is intended more as a review of
the subject, the principal object in view being to bring together what is known
of the habits of these insects to show
the uniformity of habits in genera and
species of the same genus the world
over, and, moreover, to point out just
how little real knowledge we possess of
our own species, with the hope that it
may awaken more interest in these
insects and kindle a desire in some of
our entomologists to make some effort
toward unravelling the life history of at least a few of the common species of
his neighborhood.
If every entomologist in the United
States and Canada would, during the
year 1894, make up his mind to at least
make known the habits, development
and parasites of one or two species it
surely could be done, and then what a
vast amount of new and interesting
reading we should have next winter.
How refreshing it would be to take up
one of our entomological journals, or
an experiment station bulletin, and see
some such article in place of the old,
old story, "the canker-worm, the cod-
ling moth, the chinch-bug, or the plum
Curculio ."
The subject merits attention also
from an economic standpoint, as, with
but few exceptions, all the aculeate
Hymenoptera are of the greatest eco-
nomic importance, either as fertilizers
of plants, shrubs and trees, by transport- ing pollen from blossom to blossom, or
as destroyers of injurious insects.
In order to bring out more thoroughly
the points to which I have called atten- tion, I propose to take up seriatim the
different families, give a resume' of what is known and at the same time incorpo-
rate any new facts that may have come
under my observation.
Family I. APIDAE. As the most
specialized we may begin, therefore,
with this famil~7. Excluding Apis mel-
hfica as not indigenous and the Melli-
ponae as not extending into our fauna,
we have no less than 35 genera and 520
species belonging to this family. Of
these, the genus Bombus in structure,
social habits, and in the honey-pro-
ducing qualities of its members, is
probably more closely allied to the
true honey-bee than any other of our
bees and it may, therefore, be con-
sidered the forerunner of the honey-bee. Mr. I?. W. Putnam, in "Notes on
the habits of some species of humble-
bees" (Proc. Essex inst., vol. 4, 1864,
pp. 98-104) was one of the first of our
writers to treat of some of our species. In this paper he briefly treats of the
nesting habits of Bombus ternarks
Kirby, B. fervidus Fabr., B. vagans
Smith, B. virg-inicus Oliv, B. se$ara-
tus Cr. and B. 'pennsylvanicus De Geer.
The habits of our species agree fairly
well with the observations made upon
the European species and are briefly as
follows : the female bumble- or humble-
bee, which has hibernated in some
crevice or other secure place during
the winter, appears in early spring with the first blossoms from which it can




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February 1894.1 PSYCHE. 2 1
a 0 0 0
- -
obtain pollen and almost immediately
selects a place in which to nidificate,
forming its nest of dry grass or leaves
in some hollow in the open field ; or
more frequently appropriating the de-
serted nests of field mice, either in the open field or under old stumps or boards. Here the female constructs her recepta-
cles of a waxy or plastic material, into which she deposits her gatherings of
pollen and honey-the food-supply for
the future offspring of her colony, lay- ing her eggs directly in or upon the
pollen.
The eggs first laid produce larvae,
which spin tough cocoons wherein to
undergo their transformations and all
transform into neuters or workers,
which subsequently form the greater
part of the community and become of
the greatest importance in assisting and performing the necessary economics of
the now rapidly increasing family.
These are followed later, according to
Shuckard, by males and other produc-
tive females which are, however, smaller than the normal sized individuals ; the
normal sized males and females not ap-
pearing again until the fall, when they
mate and the cycle of their life history is completed, the impregnated females
of this last brood wandering off and
hibernating and forming the nuclei of
colonies the following spring.
The number of individuals in a nest
is variable; from a dozen or more
to over two hundred individuals have
been found in a single nest. Mr. Put-
nam states that a nest of B. ternarius
contained sixty-five cells, also a number of bunches of pollen in which there
were no eggs, thirty-five contained
young and thirty were filled with honey, having their tops covered with wax and
that this was the only instance of his
finding the honey cells closed over.
Dr. A. S. Packard in "The humble-
bees of New England and their parasites, etc." (Proc. Essex inst., vol. 4, pp.
107-140) has given some interesting
and valuable observations on the species found in New England, and consider-
able new information concerning their
parasites.
Mr. Charles Robertson in "Notes on
Bombus" (Ent. news, vol. I (18go),
p. 39) and Mr. Frederick V. Covillein
"Notes on bumble-bees (Proc. ent. soc.
Wash., vol. I (1890), p. 197) from
personal observation's carried on inde-
pendently, both reached the conclusion
that Apathus elatus, a supposed in-
qiyline of Bombus fervik was in
reality the 8 of Bombus american-
o m Fabr. or B. borealis Kirby, a
species that was long confounded with
B. f ervidus.
Mr. Robertson further remarks that
Walsh in discussing the effect of mim-
icry (Proc. ent. soc. Phil., vol. 3, p.
247) mentions having once found B.
fervidus $ , surmounted by Apathus
elatus 8, and cited this as a case in
which a Bombus mistook an Apathus
for one of its own species, but remarks
^that the mistake here was on the part
of the entomologist and not on the Bom-
bus, as he had no doubt taken the true
sexes of B. fervidus."
In some particulars, Mr. Coville's
observations on Bombus borealis as in-
dicatins" a slight divergence in habits




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22 p6' 2TXL!3. [February 1894.
and thus more closely resembling the
hive-bee, warrants me in quoting some-
what largely from his very readable
paper. He says :-
The nest, originally that of a mouse, was made of dead grass and lined with wax.
It
contained when captured the queen and a
large number of workers of various sizes, as well as eggs, and larvae in various stages of development.
The precise functions of the
different sized workers were not evident, but in general the larger ones attended to the mending of the grass covering of the
nest
and to the bringing in of honey, while the smaller ones for the most part did the inside "house-work," the wax-patching and the
nursing, described below. The nursing, in- deed, was never done so far as was observed, by a large or even a medium-sized bee.
The eggs are IaiA several together, in
cavities in a mass ofwax. This is in direct opposition to the statement of Putnam (1. c.) and of various English writers consulted by me, they stating that the eggs are laid in a mass of pollen, upon which the larvae, when hatched, feed. The substance was tested first by the application of heat, when it melted precisely like bees-wax. It would not dis- solve in water, while pollen and an artificial mixture of pollen and honey readily did so. A microscopic examination of the wax showed, however, that it contained a great number of pollen grains; but this would be expected when it is considered how much pollen is used about the nest. The larvae, after hatch- ing, remained incased in a shell of wax, and soon became separated by a wall of the same substance each from its neighbor.
Their method of obtaining nourishment -
instead of by eating away the pollen walls, in which they are supposed to be incased, the workers constantly adding more to the out- side - is strikingly different. They are fed by a mixture of pollen and honey supplied to them by a worker. The operation will be
described later.
The larvae, when grown, spin a silken
cocoon, and at the end of the nymphal stage, the duration of which was unfortunately not noted, emerge by gnawing about the apex of the cocoon so as to form a lid. When the adults first come out their subsequently yellow hairs are pale, almost white. As soon as the bee has left its nymphal quarters the other workers cut away about the upper half of the cell and remove the debris. The part which is left furnishes a receptacle for the raw honey and pollen as it is brought into the nest.
When returning from the field the bees
settled down upon the alighting-block at the entrance of the box, when full laden, with a low, abruptly ceasing hum, always distin- guishable from that of a bee without honey or pollen. The bees went directly, in a most business-like way, to the pots, deposited their loads, and went away again or busied them- selves about the nest. If honey-laden, the bee perched herself on the margin of a honey- pot, lowered her head into it, and then drew her abdomen far in, thus forcing the honey from her mouth. If pollen-laden, the bee balanced herself,- with her middle and
cephalic pairs of legs, on the edge of a pollen- pot, head outward, spread her wings, and then scraped the pollen-masses from her
corbiculse by rubbing the posterior legs together.
The mode of feeding the larvae is as
follows : One of the smaller workers, which may be called a nurse-bee, goes to a honey- pot, from which she presumably draws a
small amount of honey, and proceeds next to a pollen-pot. She remains here, with her head in the pot, undoubtedly preparing a mixture of pollen and honey, for ordinarily about ten minutes.
Then going to one of
the larvae, which lie in circular form in their chambers, she injects into the cell, through a small opening previously made, usually by another worker, a brownish fluid of the con- sistency of honey. This is greedily eaten by the larva. Whether the larvae of both females and workers are fed in the same manner and



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February 1894.1 PSYCHE. 23
with the same mixture could not be decided, but from the analogous case of the honey- bee, it is to be expected that the kind of food does influence the size and function of the bee. The males, it may be added, are com- monly supposed to have come from eggs laid by the sterile females (workers).
In early August females (queens) and
males began to emerge. Both left the nest within a few days, and did not return, nor were they seen to copulate.
In the first chilly afternoon of autumn the workers become stiffened with cold, and do not return; and after a few freezing days the old queen, too, succumbs. The males also perish, and only the young queens survive the winter,
This genus is parasitized by Diptera
belonging to the genera Volucella, Con-
ops, Tachina, Coleoptera belonging to
the genera Meloe and Stylops, while
Anobiurn $aniceurn and Antherofha-
gus ochraceus Say, prey upon the
pollen stored up in their cells. A
Lepidopterous larva Ne$ho$/eryx ed"
mandsii is also supposed to be parasitic, and it itself is preyed upon by A$an/eZes ne$ho^pterygis Pack.
The genus Apathus structurally
closely resembles Bombus and the species are found living in the nests of the latter. The species are stated to be inquilinous or guest-flies, and not true parasites, and this is apparently the sum total of our
knowledge. In all the literature at my
command, I can find no direct observa-
tion respecting the rearing and develop- ment of a single species. Shuckard
makes this general statement : -
Both sexes appear to have free in and
egress to the nests of those Bombi which they infest, without any let or hindrance on the part of the latter, with whom they seem to dwell in perfect amity.
In the times of
their appearance they closely resemble the Halicti and the neighboring Bombi. Thus
the females, after impregnation in the
autumn, having hibernated during the winter in selected receptacles, come out with the first gleams of spring conjunctively with the large maternal Bornbi, in whose nests they have taken their long repose in perfect tor- pidity; and as soon as these begin to accu- mulate the masses of conglomerated honey and pollen whereon to deposit their eggs, the parasite takes advantage of it, lays hei- eggs too, and thus secures food for her
offspring.
The genus Xylocopa comprises some
of the largest bees known, many of
which closely resemble the bumble-
bees. The species are not rare and
from their method of boring into posts
and rafters, in which they construct
their nests, they are known as carpenter- bees. About a dozen species are found
in the United States.
Our most common species in the
eastern, southern and middle States
is Xylocoja virginica Drury, and its
nest is readily found in the rafters or
frame work of any old house, barn or
out-house built of soft white pine.
I have frequently found their nests
made in the railings of a porch, in posts, in rafters, in doors, in palings of fences in door frames, in window sills, etc.
Dr. Packard in his Guide, p. 132, has
given an excellent account of the nest-
ing habits of this species, as observed
by Mr. James Angus, of West Farms,
N. Y.
The species bores a cylindric& hole,
about half an inch in diameter until
the depth of ten, twelve or more inches




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24 PLS~'-CHE. [February 1894.
is attained.
At the bottom of this long
tunnel or gallery, the female now de-
posits a ball of pollen-paste in which she lays a single egg. This is then care-
fully covered over with a thin partition formed of sawdust and a glutinous sub-
stance or secretion and this constitutes the first cell. Upon this another ball
of pollen-paste and an egg is laid and
again enclosed by a partition and so
on until a series of cells, one above
another, is formed and the tunnel is
filled. The imagos hatch out in July
and August and hibernate in the middle
States during the winter months.
Mr. L. 0. Howard, in "Notes on
the hibernation of carpenter bees" (Pi-oc. ent. soc. Wash., vol. 2, 1892, p. 331)'
records having received in February a
pine branch burrowed by this species
containing living bees.
Mr. H. G. Hubbard in same publica-
tion also records Lome interesting ob-
servations made on carpenter bees in
Florida, which agreed with the writer's
own observations.
He had found in February the eggs and
the young, in various stages of development, in burrows, and in March the adult bees
ready to issue from the burrows. By April most of these had escaped and another gen-