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1928] Smith: Biology of Tapinoma 323 temporary parasite

on the odorous house ant, like its cogener, B. atlantis Forel, of Tunis
is on Tapinoma nigerrimum Nyl. The female of B. atlantis Forel after
descending from her nuptial flight is pulled into the nest of T.
nigerrimum Nyl., by the workers of that species. She soon crawls on
the back of the nigerrimum female and decapitates her. After the
death of the nigerrimum female, the workers of that species accept
the B. atlantis female and rear the brood she produces. Eventually
all the nigerrimum workers die and the colony then becomes a pure
one of B. atlantis. RELATION TO MYRMECOPHILES AND OTHER
NEST INMATES. In the nests of the odorous house ant, Mann (1911)
has found the little cricket, Myrmecophila oregonensis Bruner, and
also a little wingless wasp, Isobrachium myrmecophilum Ashm.
"Ashmead states that the genus (Isobrachium) is parasitic upon the
ants or other myrmecophilous Coleoptera," according to Mann who
writes further, "the latter being so rare in the nests of Tapinoma, it is
probable that Isobrachium is a parasite on the ant itself." The
crickets, M. manni Schimmer and nebracensis Lugger, have also
been taken in the nests of this ant. The following species of
Coleoptera, most of which are rove beetles (Staphylinidce) , have
also been taken from the nests of the odorous house ant by Mann,
Wheeler, and others: Zyras tapinomatis Mann, Myrmoecia lugubris
Casey, Nototaphra lauta Casey, Connophron longipenne Casey, and
Myrmedonia sp. The writer has found in the nests of the odorous
house ant an unidentified species of spring tail (Collembola),
termites (Isoptera), and sow bugs, Armadillidium vulgare (Latr.). In
the laboratory a species of book lice thought to be Troctes
divinatoria (Muller) appeared in the cages and fed on the' dead
bodies of the ants and other refuse. Neither were the ants observed
to attack the lice nor the lice to trouble the ants. RELATION TO
PREDATORS. The only animals known to feed on the odorous house
ants are birds and toads. - A winged male of this ant was taken from
the stomach of a toad, Bufo sp., at Anna, Illinois, by a member of
the Illinois State Natural History Survey. The following list of the
species of birds known to feed on the
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324 Annals Entomological Society of America [Vol. XXI,

odorous house ant was sent to the writer by the United States
Bureau of Biological Survey: the pigeon hawk, Falco columbarius
Linn; the American magpie, Pica pica hudsonia (Sabine) ; the
Bartramian sandpiper, Bartramia longicauda Bechet; the chimney
swift, Chaetura pelagica Linn. ; the crow, Corvus brachyrhynchos
brachyrhynchos Brehm; and the red shafted flicker, Colaptes cafer
collaris (Vigors). Bryant (1914) mentions that the western
meadowlark, Sturnella neglecta neglecta Audubon, feeds on this
species of ant and the writer has observed the English sparrow,
Passus domesticus Linn, doing the same thing. RELATION TO
PLANTS. The odorous house ant not only visits plants for the
purpose of attending insects but for the purpose of visiting also the
floral and extrafloral nectaries and other glandular excretions. The
workers have very commonly been seen lapping up exudates on the
buds of peonies. This habit apparently causes the buds to dry out to
such an extent that the flowers developing therefrom are smaller
and in many cases malformed. Dietz (1926) states that the black
lawn ant, Formica fusca var. subsericea Say, can spread peony bud
wilt by means of the workers crawling from infected to non-infected
buds, while visiting the buds for the much sought exudates. If this
be true in case of the lawn ant, it should also apply for the odorous
house ant, for it, like the lawn ant, is a persistent visitor of peony
buds. Davis of New York (1922) states that he has seen workers of
this species visiting the glands at the base of the leaves of Populus
grandidentata Michx. The writer has seen this ant in attendance on
the glands of castor bean plants. During 1926 the writer found a
fungus, Laboulbenia formicarium Thaxter, infecting several species of
ants in Urbana, Illinois, namely: Formica fusca var. argentea
Wheeler, F. neogagates Emery; Lasius niger var. neoniger Emery, and
F. pallide fulva schaufussi var. incerta Emery. Although the odorous
house ant was present on the same blocks as the species mentioned
above, this ant was not found to be infested with the fungus. In the
laboratory, colonies of the odorous house ant were 'often choked out
by luxuriant growths of the bread mold, Rhizopus nigricans Ehrh.,
and a species of Aspergillus. These molds arose from foods that
were allowed to remain in the
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1928] Smith: Biology of Tapinoma 325 cages too long,

thriving best where there was considerable excess moisture. The
writer did not find the ants outdoors to be affected by any sort of
fungus and he doubts if they ever are. FOOD. The food of the
odorous house ant under normal conditions is largely honey-dew,
supplemented by the flesh of organisms and the juices of fruits. The
ants are also very fond of the floral and extrafloral secretions of
plants. In the laboratory cages the ants occasionally fed on their
brood. It is presumed that under outdoor conditions this seldom, if
ever, happens. When the ants enter houses they are almost
omnivorous but seem to show a slight preference for sweets. In a
number of homes in Urbana, Illinois, the ants have been known to
cut through paraffine in order to reach jelly and preserves in
containers. They have been noted to infest the following foods:
honey, sugar, preserves, pies, custards, marmalades, cooked and
uncooked beef, fish, raw and fried liver, boiled and mashed potatoes,
stewed prunes, cheese, milk, ice cream, and ripe fruits. SIZE OF
COLONIES. The colonies of this ant show a wide variation in size,
perhaps due to their age. The smallest colony observed contained
only four dealated females and about one hundred workers. The
largest colony noted was estimated to contain about ten thousand
individuals, including brood and adults. An average colony may be
expected to contain between two thousand and five thousand
specimens. One of the most striking characteristics of the colonies of
this ant is the unusually large number of dealate females, all of
which no doubt take part in reproduction. The females, unlike those
of the honey bee, are most amiable toward one another. From
observations that have been made it is believed that the sexed-
forms are not produced in any but the older and stronger colonies.
Although many nests have been examined sexed-forms have been
found only in the larger and more flourishing colonies, which were
very probably from four to five years of age (Table 3).
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326 Annals Entomological Society of America [Vol. XXI,

GENERAL HABITS OF THE WORKERS. The workers of the odorous
house ant are strikingly slender and graceful in appearance. When
alarmed they run with a rapid, erratic, jerky sort of pace and
oftentimes with the caudal end of their abdomen slightly elevated.
When a nest is disturbed it is only a few seconds until the ants are
running everywhere in a somewhat helter-skelter manner. The
workers are splendid foragers, who seek food by night as well as by
day. Very seldom are they seen foraging singly but are ordinarily
observed trailing along in a file from their TABLE III. The Size of the
Colonies and the Types of Castes and Immature Stages in the Nest
at Various Periods During the Year. Total No. Dealate Male Female
Worker Date Individuals Males Females Pupae Pupa? Pupae Larvas
Eggs Apr. 1-25 104 0 4 0— 0 0 X 0 Apr. 8-26 . p 0 X 0 0 0 X 0 Apr.
12-26 p 0 X 0 0 0 X 0 Apr. 25-26 ? 0 X 0 0 0 X X June 1 1-25 ? 0 15
0 0 X X X June 6-26 ? X X X X X X X June 27-25 6,000-8,000 X *x X
X X X X Aug. 22-25 3,000-4,000 0 X 0 0 X X X Sept. 5-25 10,000 0
240 0 0 X X X Nov. 10-25 ? 0 X 0 0 X X X x — Form present. * Alate
and Dealate Females. NOTE. — Workers are not mentioned in Table
because they are present all the year and comprise the largest
percentage of individuals in the nest. nest to the source of food
supply. The ants seldom have to travel over thirty to fifty feet to find
food, and for this reason their trails are generally not long. Their
foraging activities take them into every conceivable sort of place,
such as garbage cans, commodes, dirty linens, pantries, sinks,
refrigerators, and other places too numerous to mention. Like their
cogeners, the Argentine ants, the odorous house ants are prying
little busybodies, eternally poking their antennas into everything.
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1928] Smith: Biology of Tapinoma 327 FEEDING HABITS OF

THE WORKERS. When hungry the ants eat very greedily. The writer
found that a worker will feed from three to five minutes before she
seems satisfied. During this time her gaster gradually enlarged until
the chitinous segments stood out like small islands between the
intersegmental membranes. Workers have often been observed
regurgitating food to the female or to other workers. They also feed
their larvae in this manner but never with solid particles as do some
of the primitive ants. NURSING DUTIES OF THE WORKERS. The
workers are good nurses as well as foragers. Whether there is any
particular division of labor among them, the writer does not know,
since the individuals are practically all the same size (monomorphic)
and indistinguishable. Recently emerged individuals (callows) show
considerable interest in the brood and will attempt to pick up the
brood and carry it away when the nest is disturbed. The dealated
females seem to- show little interest in the brood, leaving their care
to the workers, when these are present in sufficient numbers. The
workers show considerable attention to the female and to the brood,
around which they cluster. Since the pupae of this ant has no cocoon
covering it, the assistance of the worker for its removal at
emergence is not necessary. The writer has, however, noted a
worker removing the meconium from a larva which was preparing to
enter the prepupal stage. Workers are continually seen licking the
brood, the females, or other workers. ACKNOWLEDGMENTS. The
writer wishes to express his gratitude to Dr. C. L. Metcalf for his
unfailing interest and guidance in the work and in the preparation of
this paper. He is also grateful to Dr. W. P. Hayes, Dr. R. D. Glasgow,
and Dr. W. V. Balduf for their valuable aid and many helpful
suggestions. The writer is also indebted to the United States Bureau
-of Biological Survey and the Illinois State Natural History Survey for
furnishing valuable data, and to Dr. W. E. Britton, Mr. J. R. Horton,
and Mr. S. A. Rohwer for the loan of specimens. Mr. P. W. Mason
kindly determined the plant lice for the author.
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328 Annals Entomological Society of America [Vol. XXI,

SUMMARY. 1. Tapinoma sessile Say, a native North American ant, is
of considerable economic importance as a house pest not only in
Urbana, Illinois, but in localities in California, Nevada, District of
Columbia, Maryland, and Mississippi. 2. The synonymy and a
redescription of the species is given in this paper. The immature
stages of the ant are also described. 3. The common name, the
odorous house ant, is applied to the species for the first time. 4. The
ants overwinter outdoors as dealate females, workers, and larvae.
Workers begin foraging as early as March 7th. Egg laying and the
uniform development of the brood begins in late April and continues
until cold weather in the fall or approximately November 1st. The
overwintering larvas attain maturity in April, having required from six
to seven months to attain maturity. From April through June workers
can be produced in from five to nine weeks, whereas during the
period July through September they require only six to seven weeks
to attain maturity. Four to five generations of workers a year are
postulated. 5. Alate females were observed at various dates ranging
from June 17th to early July. Males appeared from June 10th to July
9th. Only large and strong colonies seem capable of producing
sexed-forms. Mating probably takes place both in and outside of the
nest. 6. Colonies range from a hundred to ten thousand individuals.
The average colony contains from two thousand to five thousand
individuals, many of which are dealated females; one colony
contained' over two hundred dealated females. 7. Some of the
workers lay eggs. The brood reared from such eggs attained the
prepupal stage before they were eaten by the workers. It is believed
that if they had attained maturity the adults would have been
workers. Development of -brood from worker eggs apparently takes
longer than the development of brood from female eggs. 8. The ants
show a wide diversity in their nesting habits. Nests have been found
in houses; under the bark of logs and stumps; in galls on plants; in
stems of plants; under debris; and in the soil.
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1928] Smith: Biology of Tapinoma 329 9. The species is

apparently eurythermal in disposition as it is found throughout most
of North America from the sands of the seashore to heights of over
ten thousand feet and from boggy locations to perfectly dry inland
spots. Workers have been seen foraging at temperatures as low as
fifty degrees Fahrenheit. 10. The natural food of the ants is honey-
dew, supplemented by the flesh of organisms. In houses they feed
on fruits, vegetables, meats and sweets, but seem to show a
preference for sweets. 11. A list of the species of plant lice, scale
insects, mealybugs, tree hopper, etc., which they are known to
attend is given in this paper. 12. A number of beetles, wasps,
crickets, spring tails, termites, book lice and other insects have been
found living in association with the ants. 13. A species of toad and
eight species of birds are known to feed on the ants.
BIBLIOGRAPHY. Bryant, H. C. 1914. Univ. of Cal. Pub. Zool., 11: 21-
24. Davis, W. T. 1922. Bull. Bklyn. Ent. Soc., 17:2. Dietz, H. F. 1926.
Insect Pest Survey, 6: 122-123. Doten, S. B. 1910. Nev. Agr. Expt.
Sta. Bull., 73.5: 43-44. Emery, C. 1912. Genera Insectorum, 137: 40.
Essig, E. O. 1926. Insects of Western North America. McMillan Co.,
N. Y., pp. 863-864. King, G. B. 1896. Entomological News, 7: 169.
Mann, W. M. 1911. Psyche, 18: 29. Reichenbach, H. 1902. Biol.
Centrabl., 22: 461-465. Say, Thos. 1836. Boston Jour. Nat. Hist., 1:
287. Tanner, J. E. 1892. Trinidad Field Naturalist's Club, 1: 123-127.
Wheeler, W. M. 1910. Ants, Their Structure, Development and
Behaviour. Columbia Univ. Press, N. Y., pp. 45. 1913. Bull. Mus.
Comp. Zool., Harvard, 54: 497. 1915. Bull. Amer. Mus. Nat. Hist., 34:
417-418. Woodworth, C. W. 1919. Univ. Cal. Publ., 207: 70-71.
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ANNALS E. S. A. VOL. XXI, PLATE XVIII. Marion R. Smith

Tapinoma sessile Say. Fig. 1. Worker, greatly enlarged. Fig. 2.
Dealated female, greatly enlarged. Fig. 3. Alate male, greatly
enlarged. 330