Notes on the habits of European and North American Cucujidae (sens. auct.)

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ZOOLOGICA

SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK .ZOOLOGICAL SOCIETY

FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA

VOLUME III, NUMBER 5

(Tropical Research Station Contribution Number 99)

NOTES ON THE HABITS OF EUROPEAN AND
NORTH AMERICAN CUCUJIDAE (sens. auct.)

By WILLIAM MoRTON WHEELER

12 (Us 18) hy IES) Jeb dd) 1D) ey ae Per 1D) 5 OsC 1. BT y
ZOOM OTG CoAT PeAUR KC Ni Wi 7 ORK

DECEMBER 24, 1921

Volume III, Number 5

NOTES ON THE HABITS OF EUROPEAN AND
NORTH AMERICAN CUCUJIDAE sens. auct.)

By WILLIAM MORTON WHEELER

Family Silvanidz Boving.

Oryzephilus surinamensis L. (Fig. 7.) The “saw-toothed
grain-beetle” the most abundant, most widely distributed and
best-known species of the family.’ It is cosmopolitan and gre-
garious; living in nearly all stored human foods of vegetable
origin; cereals (rice, wheat, maize, barley, etc.), ground or un-
ground or in the form of paste (macaroni), bread etc.; dried
fruits, nuts, copra, more rarely in sugar, starch, drugs, tobacco,
snuff or dried meats. The larva, which is very active, is also
gregarious, living with the beetles and evidently capable of
thriving on most of the substances mentioned (Glover, 1869;
Guillebeau, 1890; Chittenden, 1895, 1897, 1911; J. B. Smith,
1909; Girault, 1912). When ready to pupate it may make a
rude cocoon by agglutinating particles of food detritus with an
oral secretion. The pupa is sometimes free, however, i. e.,
not inclosed in a cocoon, and is attached by its hind end to
the shrivelled larval skin which has been previously attached
to the substratum (Blisson, 1849; Coquerel, 1849; Chittenden,
1895). During the summer the whole life-cycle requires about
twenty-four days, in spring from six to ten weeks. There are
six or seven generations a year in the latitude of Maryland, and
in that latitude it winters over as an adult (Chittenden, 1895).
It seems to be present wherever the Indian-meal moth (Plodia
interpunctella) is found (Chittenden, 1897) and has often been
found living with another common grain-pest, the Curculionid
Calandra oryzz (Perris, 1853; Ganglbaur, 1899). In England
and Scotland the beetle has been repeatedly taken out of doors

*I find that this or one of the closely allied species is figured by Redi
(1671) who refers to it as “vermiculus qui condita arrodit” and “vermis
conditorum et pharmacorum.” Redi also figures the grain weevil, Calandra
oryzae.

174 Zoologica: N. Y. Zoological Society [TL s'5

under the bark of trees (Fowler, 1889; Champion, 1896). “As
an instance of unusual trouble caused by this insect may be
mentioned the case cited by Taschenberg of the beetles having
invaded sleeping apartments adjoining a brewery where stores
were kept and annoying the sleepers at night by nipping them
in their beds” (Howard and Marlatt, 1896).

Oryzaephilus mercator Fauvel—The “merchant grain-
beetle,” very similar to the preceding, cosmopolitan, and recorded
as living in and feeding on pea-nuts, English walnuts, wheat,
corn-meal, cereipo fruit (Myrospermuim frutescens), candle-nuts
(Aleurites moluccana), and dried currants (Guillebeau, 1890;
Chittenden, 1897, 1911). It has been taken under the bark of
plane-trees in France, in the neighborhood of mills (Guillebeau).
Champion (1896) records it as occurring in shipments of pea-
nuts at Rouen in company with another grain-beetle, the Tene-
brionid Palorus subdepressus Woll. “The close relationship of
mercator and surinamensis makes reasonably certain their
identity as regards development, nor is it probable that they
differ in any degree in food-habits” (Chittenden, 1896).

Oryzaephilus bicornis Erich.—Cosmopolitan, but less widely
distributed and more southern than surinamensis. Lives and
breeds in wheat and dried figs in France; also taken in rubbish
at the base of fig-trees (Guillebeau, 1890; Chittenden, 1897).

Oryzaephilus gossypii Chitt—Cosmopolitan; breeding in
cotton-seed (Chittenden, 1897).

Silvanus bidentatus Fabr—Europe and United States (N.
J., Ind., Conn., So. Cala., Fla.). Under chestnut bark in the
United States (Glover, 1869). Recorded as occurring under
bark of oaks and firs in England (Fowler, 1889), of firs in
Germany (Reitter, 1911), of poplars, elms and figs in France
(Picard, 1919). J. B. Smith (1909) records it as occurring
under bark throughout New Jersey, as not rare, and as taken
most of the season. According to Picard it has the same mode
of life as the species of Laemophloeus (q. v.).

Silvanus gemellatus Duy.—Cuba and Southeastern States to
New York. Lives and breeds in maize in the field as well as in
granaries, also in wheat and over-ripe or dried fruits. It nearly

1921] Wheeler: Habits of Cucujidae 175

always first destroys the germ of the kernel and hence causes
considerable injury to seed maize in the Southern States. “It is
essentially an out of doors species, but when conditions favor its
increase may become a serious pest in granaries, as it is capable
of breeding from egg to adult in the short period of three weeks’’
(Chittenden, 1897, 1911).

Silvanus unidentatus Fabr.—Europe. Perris (1853, 1876)
states that the larva is common in France under the bark of oak,
poplar, chestnut and willow, rarer under pine bark. It is agile
and photophobie and lives gregariously with the adult beetles
among the detritus left by the larve of Bostrychus, Cerambycids
and Buprestids. In Germany under elm bark (Kaltenbach,
1874) ; in Britain under bark of beech, oak, horn-beam, etc.
(Fowler, 1889), of deciduous trees in general (Reitter, 1911).

Silvanus fagi Guérin—Europe. Under beech bark; in
England in dead branches of fir (Fowler, 1889); in Germany
under fir bark and fir-cones (Reitter, 1911).

Silvanus planatus Germ.—Eastern United States (N. J.,
Ind., Fla.). Under pine-bark in New Jersey (J.B. Smith, 1909).

Cathartus advena Waltl.—Cosmopolitan. Lives and breeds
in stored wheat, rice, corn in stack, grain, meal, middlings, flour,
dates, figs, lichi nuts, table beans, cacao-beans, edible tubers, etc.,
but apparently only when these substances are not kept dry and
clean. “In breeding experiments recently conducted by the writer
it failed to develop in fresh grain or meal, but bred freely in
corn-meal which was moistened and produced mold. The beetles
particularly fed freely on the molds, of which there were three
or four species, and it would appear that this is the normal habit
of the insect”’ (Chittenden, 1897). Pierce (1917) cites C. advena
among the insects infesting teak in India, stating that it “breeds
in leaves, forming galls, causing leaves to drop” (sic!). Perris
(1876) found the larva in lichi nuts in company with larve of
O. surinamensis. Fowler (1889) states that C. advena is some-
times found out-of-doors in England and that Waterhouse took
it under cut grass. In New Jersey it is ‘rare under bark; more
common in stored grain, fruits, nuts, ete., particularly such as are
spoiled” (J. B. Smith, 1909)

176 Zoologica: N. Y. Zoological Society LU 55

Cathartus cassizx Reiche-—Cuba and Southern States.
Glover (1869) describes the larva as feeding on maize kernels,
near the germ, and also on the exposed seeds of cotton bolls.
Doran (1892) found this beetle breeding in bran and middlings
and producing a temperature 42 deg. F. higher than that of the
room in which it was living!

Cathartus longulus Blatchley.—Indiana. “Sifted from bor-
ders of Sphagnum marsh” (Blatchley, 1910).

Cathartus excisus Reitter—Taken in Germany in Havana
tobacco (Ganglbaur, 1899; Reitter, 1911).

Cathartus quadricollis Guérin—Imported into Germany in
Havana tobacco and also in the fruits of Cassia fistula (Gangl-
baur, 1899) ; imported in Havana cigars (Reitter, 1911).

Nausibius clavicornis Kug.—Cosmopolitan. “Under bark
and in sugar” (Glover, 1869). ‘“‘Often found on ships in plant
wares. It also lives in old nests of South American bees” (Gangl-
baur, 1899). Introduced into Britain, found in sugar, etc.
(Fowler, 1889). In New Jersey under bark and also occasionally
in store-houses (J. B. Smith, 1909). In Indiana “two specimens
were taken with a dozen other species at sap beneath the bark
of a soft maple tree. Leconte states that it occurs usually in
rice, sugar and other articles of commerce throughout the United
States” (Blatchley, 1910).

Telephanus velox Haldem.—United States (N. J., Ia., Ind.,
Conn.). “Very common under rubbish of all sorts and as its
name implies, runs with remarkable swiftness” (Casey, 1884).
“Occurs more often beneath stones, chunks and dead leaves than
beneath bark. When exposed it usually remains quiescent with
antenne folded against sides; but if touched it runs with great
swiftness, whence its specific name’ (Blatchley, 1910).
“Throughout the State (N. J.) under stones and old leaves;
rarely under bark; may be sifted out from fall to late spring”
(J. B. Smith, 1909).

Family Cucujide (Boving emend.)

Cucujus clavipes Fabr.—Atlantic States to Illinois and Iowa;
a beautiful scarlet red insect, represented in the Pacific States

1921] Wheeler: Habits of Cucujidae ial

by the var. puniceus Mann. The larva is said to be predatory
(Le Baron, 1874; J. B. Smith, 1909). Le Baron (1874) figures
the larva; Dimmock (1884) records the adult beetle as occurring
under decaying butter-nut bark. Hamilton (1886) gives the
following account of the insect: ‘“‘The very depressed form of
this well known beetle indicates, a priori, its subcortical habit,
and no other has power to adapt its tastes to a greater variety
of timber—locust, maple, oak, hickory, gum, buckeye, etc.—are
all alike to it. The larve do not eat the wood nor the bark,
living apparently on the moisture existing between the two. They
are elongate, much depressed, brownish yellow, and scarcely to
be distinguished from those of Dendroides canadensis. Some-
time in September, the larva having matured, constructs a circu-
lar cell from small particles of the decaying bark and wood, and
in this completes its transformations before severe frost, but the
beetle does not quit the cell till the following spring. I have
never known any of these insects to be taken elsewhere than
under bark, though they undoubtedly fly, being possessed of a
good pair of wings. On the 10th of October, fifteen newly dis-
closed individuals and several pupe# were taken under the bark
of a gum log; the latter are depressed like the beetle, pale at first,
the eyes, antenne and portions of the legs gradually changing
to black, and the elytra becoming red after disclosure. This
insect is annual.”

Cucujus haematodes Erich—Europe. In Germany under
bark of maples, rarer under bark of fir-stumps (Reitter, 1911).

Cucujus cinnaberinus Scop.—Europe. Habits like those of
haematodes (Reitter, 1911).

Dendrophagus crenatus Payk.—Europe. In Germany under
oak bark; gregarious (Kaltenbach, 1874) ; in Britain under bark
of fir (Pinus sylvestris), less frequently under bark of larch
(Larix europxa) (White, 1872; Fowler, 1889). According to
White the larva has been supposed to be carnivorous, but is truly
phytophagous, feeding on the decaying inner bark of dead and
usually prostrate trees of the species mentioned. The beetles are
agile and seem to come out at night and run about on the bark.
The eggs seem to be laid in the spring by hibernating females;
the larve feed twelve to fourteen months, becoming pupz during

178 Zoologica: N. Y. Zoological Society (ust

the second summer, the beetles emerging in August. The larva
is very quick and agile and when disturbed moves the hinder part
of its body quickly from side to side. When about to pupate “‘it
attaches itself firmly to a piece of bark by the thirteenth segment,
and the pupa remains attached by its anal segment to the larva-
skin.” Perris (1876) believes that White is mistaken in his
account of the food-habits of the insect, and that it is carnivorous
or coprophagous, like Brontes planatus (q.v.).

Emporius signatus Frauenf.—Imported into Germany from
the tropics in Havana cigars (Ganglbaur, 1899).

Pediacus depressus Herbst—Recorded from Europe, Van-
eouver and the United States (So. Cala., Mich., Lake Superior,
Colo., Pa., S. C.). In Britain under oak-bark, in mold-like fungus ;
also in stores on board a yacht (Fowler, 1899). Under bark of
deciduous trees in Germany (Reitter, 1911).

Pediacus fuscus Erich—Europe and North America, Alaska,
Labrador, British America, Mich., Neb., Colo., N. Mex., Lake
Superior, and probably cosmopolitan. In Germany under pine
and oak bark (Reitter, 1911), and under the bark of firs (Kalten-
bach, 1874). In Britain under bark and in chunks of freshly
cut oak, beech and horn-beam (Fowler, 1889).

Brontes planatus L.—Europe. Perris (1853) describes the
larva as livid reddish, punctate with carmine dots, and as being
very agile and supple. In France it is common throughout most
of the year under the bark of various trees, but most often
under oak or pine bark. Perris believes the larva to be carni-
vorous, because it is always found among the larve of Tomicus
or with Poduri or mites ‘dont elle fait, sans doute, sa nourri-
ture.” ‘When about to pupate it attaches itself to a flat surface
by means of the tubercle of its last segment, then the skin splits
along the back and is pushed back to the posterior end of the
body, where it remains much wrinkled.” In Germany under
bark of deciduous trees, especially of oaks (Kaltenbach, 1874;
Reitter, 1911); in Britain under bark of dead beech trees
(Fowler, 1889). According to Perris (1876) the larva is
predaceous and a scavenger, living under the bark of various
trees, including, chestnut, especially where the larve of other

1921] Wheeler: Habits of Cucujidae a9

beetles have lived. All the stages are passed through in several
months, the young larve being found in the spring, the adults
in late summer. The latter hibernate under the bark. According
to Ganglbaur (1899) the larva is “undoubtedly carnivorous.”

Psammoechus bipunctatus Fabr—Europe. Recorded in
Britain from marshy places at roots of grass and in refuse
(Fowler, 1889) ; in Germany on the banks of streams or ponds
under dead rushes (Ganglbaur, 1899; Reitter, 1911).

Psammoechus desjardinsi Guérin.—Cosmopolitan (Casey,
1884) ; Florida (Leng, 1920).

Family Laemophloeide Boving.

Laemophloeus alternans Erich—Cosmopolitan; “found
everywhere” (J. B. Smith, 1909). In Germany under bark of
conifers in galleries of beetles of the genus Pityogenes (Reitter,
1911). In France occurring occasionally in the galleries of
Hypoborus ficus, a bark beetle of fig-trees, but also in galleries
of other bark beetles in other plants (Perris, 1853; Picard,
1919).

Laemophloeus ater Oliv—Europe. In France in dead or
dying stems of broom and gorse, preying on the larve of the
bark-beetle Hylesinus rhododactylus (Perris, 1853). In dead
‘stems of broom and under elm-bark in Britain (Fowler, 1889).
Rare under bark; common in bran and middlings in Germany
(Reitter, 1911). Occurs in France not only in the galleries of
Hypoborus ficus in fig-bark, but also in the galleries of other
Scolytids in other plants (Picard, 1919).

Laemophloeus biguttatus Say—United States (Ind., Neb.,
Fla., So. Cala., N. J.). Throughout the State of New Jersey
under bark; fall to mid-summer. (J. B. Smith, 1909.)

Laemophloeus bimaculatus Payk.—Europe. Under bark of
oak, beech and horn-beam in Britain (Fowler, 1889); in Ger-
many under beech bark (Reitter, 1911).

Laemophloeus castaneus Erich.—Europe. Under birch and
chestnut bark in Germany (Reitter, 1911).

180 Zoologica: N. Y. Zoological Society UE S59

Laemophloeus corticinus Erich.—Europe. Under oak-bark
in Germany (Reitter, 1911).

Laemophloeus clematidis Erich.—Europe. In Clematis
vitalba. A constant companion of Xylocleptes bispinosus, feeding
on the refuse and dejecta in its galleries (Perris, 1853; Kalten-
bach, 1874; Reitter, 1911). In dead stems of Clematis vitalba
in Britain (Fowler, 1889).

Laemophloeus denticulatus Preysl—Europe. ‘The larve
probably merely accompany the larve of really injurious
xylophagous beetles. According to Hartig they are found in
the cones of conifers, according to Ratzeburg also under dead
bark, according to Hellwig and Panzer under the bark of linden
trees” (Kaltenbach, 1874).

Laemophloeus dufouri Laboulb.—Europe. The larvee were
found by Perris (1853) under pine-bark hibernating in January
with the pupe and beetles in the galleries of Crypturgus pusillus.
He believed that the larve feed on those of the Crypturgus, but
more probably they merely eat the dejecta in the burrows.

Laemophloeus duplicatus Waltl—Europe. Under bark of
beech, oak, etc., in Britain (Fowler, 1889) ; in Germany under
beech bark (Reitter, 1911).

Laemophloeus fasciatus Mels.—United States (Ind., Pa.,
Minn.). In Minnesota it “is found under bark, but is also very
common near saw-mills, especially in those in which hardwood
is sawed. In running over persons engaged in mills it will bite,
even without provocation, and may cause bad sores’ (Lugger,
1899).

Laemophloeus ferrugineus Steph—Cosmopolitan (Casey,
1884). In Britain in hay-stack refuse, rarely under bark; also
in granaries (Fowler, 1889). In Germany in rice, bran, mid-
dlings and meal (Ganglbaur, 1899; Reitter, 1911; Chittenden,
1911). “Its larva, which does not differ from that of other species
of the genus, has been described by Carpentier (Bull. Soc. Linn.
Nord. France, April, 1877, 3 pp. 239-241). It lives under the
bark of oaks, cherry trees and, it is said, in fig-trees inhabited
by wood-boring insects, but also in grain attacked by Calandra,
in bran, flour, etc.’’ (Picard, 1919).

1921] Wheeler: Habits of Cucujidae 181

Laemophloeus hypobori Perris.—Europe. (France). Living
only in the galleries of a bark-beetle Hypoborus ficus, in fig-trees.
Both the adult beetles and the larve live gregariously and feed
on the dejections, exuvie and all kinds of detritus which they
find in the empty galleries. They occur more rarely in the
brood galleries that are still inhabited by the parent Hypobori.
Both larve and adults are found throughout the year and both
stages, but especially the adults, are found hibernating (Picard,
1919). This author gives an excellent account of the beetle and
its habits.

Laemophloeus juniperi Grouv.—Europe. Under bark of
Juniperus in galleries of the bark-beetles Phloeosinus bicolor Br.
and thujxe Perris (Reitter, 1911). ‘“‘At Monpellier this is the
rarest species in fig-trees; I know of only one capture made in
August by J. Lichtenstein. L. juniperi is nevertheless common
in Herault, as in the whole South. I often found it under the
bark of elms riddled by Pteleobius vittatus and kraatzi; it is also
found in the galleries of Hypoborus (Liparthrum) mori of the
mulberry, of Phloeosinus thujx and bicolor infesting Juniperus,
Cupressus, Thuja and other conifers, of Phloeotribus scarabaeo-
ides of the olive, etc.”’ (Picard, 1919). The larva, which is copro-
phagous and detritivorous, has been observed and described from
Thuja infested with Phloeosinus thujx by Decaux (Bull. Soc. Ent.
France, June 25, 1890, p. 125) and is redescribed by Picard.

Laemophloeus modestus Say.—United States (Tex., Fla.,
D. C., N. J.). ‘Taken in hemp-seed, the interior of which had
been entirely eaten out’ (Glover, 1869). In New Jersey
occurring under bark and in siftings (J. B. Smith, 1909):

Laemophloeus monilis Fabr.—Europe. In Germany under
beech and linden bark, gregarious in the burrows of Tachrorhy-
chus bicolor (Reitter, 1911).

Laemophloeus perrisi Grouv.—Europe. Occurring in the fig.
“Tt is also known from the Lentiscus and pines infested with
Pityophthorus. This is a Corsican insect, very rare in France,
where it has scarcely been seen, except in the Provence” (Picard,
1919).

182 Zoologica: N. Y. Zoological Society [Ss

Laemophloeus pusillus Schén—Cosmopolitan and common
(Casey, 1884). In Britain in granaries, imported with grain
(Fowler, 1889). Occurs commonly in cereals, but probably in
the main predaceous and scavengering (Chittenden, 1911). In
rice, grocery wares, etc., in Germany (Reitter, 1911).

Laemophloeus testaceus Fabr.—Europe. Similar to L. denti-
culatus but occurring under linden bark (Kaltenbach, 1874), and
under dry beech bark (Reitter, 1911). Perris (1853, 1876) found
the larve under oak bark in galleries of Tomicus and under
chestnut bark with the larve of the bark beetle Dryocoetes capro-
natus on which it was preying, but also living on the dejections
and capable of developing to maturity in the absence of the
Dryocoetes. Also found under bark of dead oaks with the larve
of the same Scolytid and in the elm in the galleries of Hylesinus
vittatus and kraatzi.

Laemophloeus turcicus Grouvy.—Cosmopolitan. Imported
ito Germany in dried fruits, especially prunes, from the orient
(Reitter, 1911).

Laemotmetus ferrugineus Gerst—Cosmopolitan. Imported
into Germany in rice (Ganglbaur, 1899; Reitter, 1911).

Laemotmetus rhizophagoides Walker.—Cosmopolitan. Found
in stored rice in Berlin, Germany (Chittenden, 1911).

Lathropus vernalis Erich.—United States (Fla., Miss., N. J.,
Conn.). Throughout New Jersey, secured by beating dead
branches, May to July (J. B. Smith, 1909).

Lathropus sepicola Miiller—EKEurope. According to Perris
(1876) the larva lives under elm-bark in the galleries of the bark
beetles Scolytus multistriatus, Hylesinus vittatus and H. kraatzi
and feeds on their dejections. The perfect insect emerges in
May. Reitter (1911) often beat the latter from dry wood
fences.

Phloeostichus denticollis W. Redtb—Europe. Under the

inner bark of maple-trees, together with the larval stages
(Weisse, 1897; Reitter, 1911).

Prostomis mandibularis Fabr—Europe and Pacific States
(Or., Nev., Cala.) and Vancouver. In very rotten, damp wood

1921] Wheeler: Habits of Cucujidae 183

of deciduous trees. The larve are often present in great numbers
with the beetles (Ganglbaur, 1899; Reitter, 1911). Perris (1876)
found the larve in rotten chestnut wood and states that Curtis,
Chapuis and Candéze took them in rotten oak wood.

Hemipeplus marginipennis Lec.—Cited only from Georgia
and Florida (Leng, 1920). According to Schwarz a rather
abundant species under palmetto bark (Casey, 1884).

Hemipeplus microphthalmus Schwarz—Cited only from
Florida (Leng, 1920), where it was taken flying at night by
Schwarz (Casey, 1884).

Inopeplus praeustus Chevr.—Antilles. Adults and larve
living gregariously in the burrows of a Scolytid in branches of
cacao. The larva is peculiar in having the prolongations of the
ninth abdominal segment in the form of forceps (de Peyerimhoff,
1908).

Family Scalidiide Boving.

Catogenus rufus Fabr.—United States (Ind., Ia., Md., N. C.,
Fla.). Fiske (1905) has shown that the larva is an external
parasite on the larve of Braconid Hymenoptera and Cerambycid
pupe; “its habits differing in no essential feature from those of
many species of external Hymenopterous parasites. The adult is
fairly common throughout the South, and is found beneath the
loose bark of recently dead and dying trees, both conifers and
deciduous. It occurs at nearly all seasons of the year, but is
especially common during the late fall and early spring, and is
found hibernating in situations similar to the above mentioned.”
Fiske seems to have overlooked a note on this beetle by Dimmock
(1884), who says: ‘In Connecticut it is common beneath the
loose bark of the trunks of hickory trees, and I have reared its
larva which fed upon a pupa of Elaphidion parallelum, a borer
in hickory.”

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QL Wheeler, William Morton.
591 Study of some social
B74W48 beetles in British Guiana,
Ent. 1921.