Transactions of the Academy of Science of St. Louis “22!
VOLUME XXV
TITLE PAGE AND INDEX
JANUARY, 1924
AUGUST, 1928
Published Under Direction of in Corea Elv ee
JUN 10 i929
PPP PD
Academy of Science of St. Louis
LIST OF OFFICERS, 1928
2 a. Toone
” -VWice-PRESIDENT .e-eceeeneeenneneensnennene seo Loeb
RECORDING GECRETARY 1: .cccccccceeceeeceeneeeeeeesee A. G. Pohlman
CoRRESPONDING SECRETARY.........-------.J- I, Shannon
‘PRESIDENT
TREASURER. H. E. Wiedemann
LIBRARIAN oe ree
Cire aii 8 ee H. C. Irish
: ) Hermann von Schrenk
CONTENTS.
TABLE OF CONTENTS
Last oF OFFICERS
Papers PUBLISHED. January, 1924, to August, 1928:
SE
W. F. Cuapp.—Three New Species of Teredo,
Plates I-II[I.—Issued April, 1924
R. Wauter Mitus.—Mediecal Fads and Fanceies.
Lf
o
—Issued April, 1924 Coke
Aueustus G. Pontman.—The Natural History
Museum Movement in St. Louis.—Issued
April, 1924
Puit Rau.—The Biology of the Roach, Blatta
orientalis Linn.—Issued April, 1924
W. F. Cuapp.—Notes on the Stenomorphie Form
of the Shipworm, Plates’ IV-V.—Issued
January, 1925
J. T. BucHHouz anp E. J. PaumMer.—Supple-
ment to the Catalogue of Arkansas Plants,
Plates VI-X1I1.—Issued June, 1926...
Put Ravu.—The Ecology ot a Sheltered Clay
Bank; a Study in Insect Sociology, Plates
XIV-XXI—Issued August, 1926.00.
CHARLES Rosertson.—Florida Flowers and In-
sects.—Issued September, 1927
Pui Rav.—Field Studies in the Behavior of the
Non-Social Wasps, Plates XXII-XXXII.—
Issued June, 1928.
16
29
121
49
166
i 4,
Transactions of the Academy of Science of St. Louis
/
Volume XXV, No. 1°
O.&
_ THREE NEW SPECIES OF TEREDO
W. F. CLAPP
THREE NEW SPECIES OF TEREDO.
W. F. Crapp.
The material from which the following descriptions
were drawn, was removed from test-blocks of wood
placed in the water by the Committee on Marine Piling
Investigations of the National Research Council. For
descriptions of the test-blocks, and also for notes regard-
ing the nomenclature used in the descriptions of the
various characters of the shell and pallets, and for ref-
erences to recent literature on the subject, consult
Clapp, 1923, Proc. Bos. Soc. Nat. Hist., vol. 37, no. 2,
pl. 3-4, p. 31-38, text fig. 1.
I am deeply indebted to Prof. S. C. Prescott of the
Massachusetts Institute of Technology, Cambridge,
Mass., for laboratory facilities and for other assistance.
; Trans. Acad. Sci. of St. Louts
TEREDO (TEREDO) PORTORICENSIS SP. NOV.
Plate I, figs. 1-7.
Shell subglobular, white, covered with a transparent,
colorless: periostracum. The juncture of the anterior
with the anterior-median area clearly marked by a
broadly curved, slightly incised line. The ventral edge
of the anterior area, forming an angle of about 100°
with the anterior edge of the anterior-median area.
Externally, the anterior area with the usual denticu-
late ridges, the ventral posterior portion, with the ridges
of about one-half the width of the intervening spaces.
Dorsally the ridges are only one-fourth as wide as the
spaces between them. Beginning at the ventral posterior
edge, there are eight of these ridges to the millimeter,
each ridge bearing approximately seventy denticles to
the millimeter (fig. 1). The anterior-median area, at a
point opposite the ventral edge of the anterior area, is
one-quarter of the width of the entire median area. The
denticulate ridges on this area, in a line continuous with
the ventral edge of the anterior area, average twenty-
five to the millimeter. In the type there are eleven of
these ridges, which bear the usual broad denticles, there
being twenty-eight denticles to the millimeter (fig. 2).
The ventral ends of these ridges can be clearly seen
continuing as sharp growth lines over the entire median
area, but becoming less distinct on the auricle. The
middle-median area is milk-white, in contrast to the
semi-transparent anterior-median area, and is separated
from the anterior-median area by a thin, narrow, trans-
parent band. The posterior-median area is milk white
and occupies one-half of the entire median area. The
_ auricle is semi-transparent, showing more or less irreg- _
ular growth lines, and the periostracum is here thicker —
than elsewhere on the shell. :
Three New Species of Teredo 3
Internally, in the left valve a short, broad, flat hinge
plate, directly beneath the umbone. The blade slightly
more than one-half the length of the shell, of nearly uni-
form width for its entire length, the lower half reflected
posteriorly. The ventral knob large. The internal shelf
of the auricle well marked.
Pallets (fig. 3) of the type of Teredo navalis Linné.
The stalk of about the same length as the blade, and
merging into it by a gradual curve. The lower half of
the blade white, the calcareous part of the upper half,
as seen through the transparent chitinous covering,
slightly cupped, the lateral portions extending farther
distally than the median. The upper half entirely com-
posed of transparent, yellowish horn-colored periostra-
eum, which is deeply cupped distally for more than half
its length, the outer surface being slightly less deeply
cupped than the inner, and with a deep, narrow sinus -
at the center,
The posterior end of the tube with two short, nar-
row, low ridges arising from opposite sides of the in-
ternal wall; these ridges continue posteriorly beyond
the shelly portion of the tube as sharp points.
The type specimen (Mus. Comp. Zodl., 45303) is from
San Juan, Porto Rico. Additional specimens from the
type locality are also in the U. S. National Museum.
The measurements of the type are:
Total length of tube 40 mm.
Shell, height 3.2 mm., length 3.1 mm.
Pallets, length 3.8 mm., divided equally between
blade and stalk, width of blade 0.8 mm.
Teredo portoricensis is more closely related to Teredo
bartschi Clapp than to any other described species. The
variation of the shell characters in each species is so
great that only very slight constant differences can be
“s Trans. Acad. Sci. of St. Louis
seen. The normal shell of a mature specimen of T.
portoricensis is smaller than that of T. bartschi. The
length of the apophysis always proportionately less.
The partitions in the tube, while always present, are
much lower and shorter than in 7. bartschi. The pallets
resemble those of 7. bartschi, but constantly differ from
them in having the blade longer and narrower, the june
ture of the blade and pallet hardly perceptible, and
the basal portion of the blade more gradually expanded.
Seen through the periostracum, the calcareous portion
of the blade of the pallet of 7. bartschi appears cone
shaped, whereas that of T. portoricensis is the opposite,
being deeply cupped at the center. In T. portoricensis
the periostracum on the outer face of the blade is less
deeply cupped than that on the inner, while in 7. bartscht
the reverse is true.
This species has been found in the test-blocks placed
by the Committee on Marine Piling Investigations, at
the following locations: Guantanamo, Cuba; San Pedro
de Macoris, Santo Domingo; Port au Prince, Haiti; San
Juan, Porto Rico; St. Thomas, Virgin Islands; Coco
Solo, Panama, and (one specimen) Key West, Florida.
At Guantanamo, Cuba, wood placed in the water on
April 10th, contained specimens 10 mm. in length on
May 10th; 60 mm. in length on June 10th, and 75 mm.
in length July 10th.
At Port au Prince, Haiti, wood submerged December
1, 1922, contained on January 1, 1923, many 5 mm. spec
imens; on January 15th, 30 mm. specimens, and, on Feb-
ruary Ist, 30 mm.:specimens with the gills well filled
with many fully developed embryos. Wood placed in
the water at this location on June 1, 1923, although well
filled with several other species of shipworms, contained
Three New Species of Teredo 5
no specimens of 7. portoricensis as late as September 3,
At San Pedro de Macoris, Santo Domingo, wood sub-
merged December 1, 1922, contained 20 mm. specimens
on February 1, 1923, many of them with well-developed
embryos.
At San Juan, Porto Riico, wood placed March 20,
1923, contained on May 30th, 7. portoricensis 40 mm.
long with embryos.
At St. Thomas, Virgin Islands, wood placed April 1,
1923, contained many-30 mm. specimens on June Ist.
At Coco Solo, Panama, wood submerged December 4,
1922, contained many 30 mm. specimens on January 19,
1923, and on February 19th, many specimens with well-
developed embryos in the gills.
It can be seen from the above records that this species
may grow to be 60 mm. in length in a period of two
months, or at the rate of approximately 1 mm. a day,
and that specimens with a total tube length of but 20
mm. may possess well-developed embryos within the
gills. Its rapid growth and early sexual maturity ren-
der it one of the species most frequently found in the
West Indies, and the destruction caused by it is con-
siderable.
Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
PLATE
(All figures reduced *%)
Teredo portoricensis
Denticulate ridges of the anterior portion. x 100.
Denticulate ridges of the anterior-median portion. x 100.
Pallets. x 14.
Exterior of right valve. x 14.
Exterior of left valve. x 14.
Interior of right valve. x 14.
TRANS. AcaAp. Scr. oF Sit. Louts, Vou. XKV PLATE )
FIG. 4
FIG. 2 FIG. 3
Three New Species of Teredo 7
TEREDO (ZOPOTEREDO) JOHNSONI* SP. NOV.
Plate II, figs, 8-15.
Shell subglobular, white, covered with a thin, nearly
transparent, colorless periostracum. The narrow incised
line separating the anterior and the anterior-median
areas very slightly curved. The ventral edge of the an-
terior area meeting the anterior edge of the anterior-
median area in a nearly straight line, forming an angle
of approximately 90°.
Externally, the anterior area large, with many evenly
spaced, denticulate ridges, which are of about the same
width as the intervening spaces. There are sixteen of
these ridges to the millimeter on the posterior-ventral
portion of this area, each ridge bearing one hundred and
twenty minute denticles to the millimeter (fig. 8). The
anterior-median area occupying, at its widest part, one-
third of the entire median area. The denticulate ridges
on this area, along a line continuous with the ventral
edge of the anterior area, average thirty to the milli-
meter. There are twenty-six of these ridges in the type
specimen, each ridge bearing approximately thirty-three
denticles to the millimeter (fig. 9). The middle-median
area is narrow, divided longitudinally into nearly equal
halves, the anterior half, with the continuation of the den-
ticulate ridges, showing as narrow, diagonally descend-
ing growth lines, which curve upward, and become more
or less obscure on the posterior half of the middle-median
area. The posterior-median area large, occupying more
than half of the entire median area, nearly smooth, show-
ing only occasional, faint, incised growth lines. The
auricle very small, being merely a continuation of the
*I take pleasure in naming this species for Mr. A. A. Johnson, As-
sistant to the Director of the Marine Piling Investigations Committee
of the National Research Council.
8 Trans. Acad. Sci. of St. Louis
posterior-median area, with no trace externally of a
separating groove or concavity.
Internally, a small, square hinge-plate in the left valve.
The blade two-thirds of the length of the entire shell,
thin, broad dorsally, the middle and ventral portions
narrower, its entire length reflected slightly posteriorly.
The ventral knob narrow, long, its base extending dor-
sally for a considerable distance. The juncture of the
anterior with the anterior-median area marked by a nar-
row, thickened chord. The juncture of the auricle with
the posterior-median area hardly visible.
Pallets (fig. 10) with long, stout, opaque white stalks.
The blade short, broad, investing a considerable por-
tion of the upper part of the stalk in a thin sheath. The
outer face convex. The proximal third of the blade eal-
eareous, arising from the stalk in an abrupt curve. The
middle third swollen, covered with a light horn-colored
periostracum. The distal third covered with a dark
chestnut colored periostracum with a deep central sinus
dividing this portion of the blade into two shallow cups,
the outer face considerably more deeply indented than
the inner. The inner face of the blade flat, its distal two-
thirds covered with a periostracum irregularly streaked
with narrow bands of light and dark chestnut.
The posterior end of the tube with a long delicate
partition (fig. 11) dividing the tube into equal parts.
The type (Mus. Comp. Zo6l., 45306) is from Guan-
tanamo, Cuba.
The measurements of the type are:
Shell, height 4.5 mm.; length 4.5 mm.
Pallets, total length 4.3 mm.; length of stalk 3 mm.;.
width of blade 1.6 mm.
The shell of this species is very closely related to that
of Teredo clapy Bartsch (Proc. Biol. Soc. Washington,
Three New Species of Teredo 9
1923, 56, p. 96). However the apophysis is shorter and
narrower, the ventral knob smaller. The entire median
area in Teredo johnsoni is always proportionately
broader than in Teredo clappi. The range of variation
of the denticulate ridges of both species is so great: that
no constant difference can be established, but the angle ~
formed at the juncture of the ridges of the anterior area
with those of the anterior-median area is always ap-
proximately 90° in Teredo johnsoni whereas, in Teredo
clapyi it is constantly obtuse, being rarely less than 100°.
The pallets are very different from those of any pre-
viously described species of Zopoteredo in that they are
more nearly truly double-cupped, being in this respect
more like Teredothyra. In Teredo somerst the charac-
teristic median sinus can be seen only in the pallets of
the very young, never persisting in the mature speci-
mens. In Teredo clapyi it is frequently obscured or lost
entirely in mature individuals, although always present
in immature specimens. In Teredo johnsoni the sinus
is constant in specimens of all ages.
Specimens of this species have been found in the test-
blocks placed by the Committee on Marine Piling Inves-
tigations at the following locations: Guantanamo, Cuba;
Port au Prince, Haiti; Fajardo and San Juan, Porto
Rico; St. Thomas, Virgin Islands.
This species is comparatively rare and the destruc-
tion caused by it very slight. Frew specimens exceed
60 mm. in total length.
At Guantanamo, Cuba, a test-block placed in the water
on October 1, 1922, contained specimens 20 mm. in length
on December 30th. A special block made of shingles,
placed in the water in April, 1923, contained several 60
mm. specimens July 7, 1923. At Port au Prince, Haiti,
10 Trans. Acad. Sct. of St. Louis
a test-block submerged June Ist and removed August
14th, contained Teredo johnsoni with 30 mm. tubes. At
San Juan, Porto Rico, a test-block placed October 1,
1922, contained 30 mm. Teredo johnsoni on January 2,
1923, and wood placed in the water July 1, 1923, con-
tained small specimens on August 1, 1923. At Fajardo,
Porto Rico, wood submerged from January 9 till April
30, 1923, contained several specimens of this species. At
St. Thomas, Virgin Islands, wood submerged four
months and removed January 1, 1923, contained sev-
eral specimens.
any
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Three New Species of Teredo
EXPLANATION OF PLATE
PLATE
(All figures reduced 3/7)
Teredo johnsoni
See AOR OT ALLELE OP
EONS
Denticulate ridges of the anterior portion. x 130.
Denticulate ridges of the anterior-median Ia x 130.
Pallets.
x
Posterior portion of the tube. x 13.
Exterior of right valve.
Exterior of left valve.
Interior of left valve.
Interior of right valve.
bal
_
13.
x 13.
= 13.
12 Trans, Acad. Sci. of St. Louts
TEREDO (ZOPOTEREDO) FULLERI* SP. NOV.
Plate III, figs. 16-22.
Shell subglobular, small, bluish white, covered with a
strong, transparent periostracum.
40 days 1 1 Total. 269 29
42 days 7 3
Thus we see that the average duration of life for the
males was 40.2 days, and for the females 43.5 days. This
difference is not great, but these few figures show at
least that the females are inherently longer lived than
the males, a condition that is often found in the insect
world.
The Biology of the Roach 63
The data gathered in 1916 indicated that July was the
natural period of death for the roaches, both those kept
for observation and for the population at large. By
August 2nd, all those in the jars had died, and simul-
taneously the adult roaches at large in the building bad
disappeared.
The housewife has a justifiable abhorrence of these
creatures when they are adult; when thousands of
nymphs infest her home, either she sees them not, or
she bothers not about them, but the adults are con-
spicuous and at once arouse her wrath. Forthwith she
hies her to the nearest chemist and gets some good
exterminator; two weeks later she will in good faith
gladly tell the merits of ‘‘X¥YZ’’ paste, and one cannot
convince her that she spread her poison just at the time
when the roaches were dying a natural death, and that
they would have disappeared at that time of the year
regardless of whether or not she spread her paste.
Number of egg cases deposited.—The female deposits
during her lifetime from one to four egg cases. Fifty-
five roaches were confined for the purpose of getting
data on oviposition. The following table presents the
data.
Egg cases. Females.
1 8
2 19
3 19
4 9
The egg cases are not dropped as soon as they are
formed, but they are slowly pushed out from the abdo-
men, and after they have been carried about for two or —
three days, they are finally dropped. I am inclined to
suspect that the duration of the period of waiting is
determined in greater part by inherent physiological
64 Trans. Acad. Sci. of St. Lous
conditions, and in a lesser part to the temporary en-
vironment, i. e., the dropping of the egg case is to some
degree postponed until a place is reached where it may
be deposited amid proper surroundings of moisture, tem-
perature, seclusion, etc.
Age of females at time of first oviposition—Records
were kept of the lapse of time between the date when the
female roach went through the final moult and became
adult, and the day when she deposited her first egg case.
These intervals were 8, 11, 11, 13, 14, 15, 18, 19, 22,
and 24 days.
Thus the normal females in this series never deposited
before they were 11 days old (the 8-day individual was
virgin), and one deposited her first egg case at the age
of 24 days. Averages, however, are of no significance
in this case, since in all probability egg laying is closely
connected with mating, and no records were kept of the
time that elapsed between mating and oviposition. This
oviposition occurred from the third week in May to the
middle of June, but no correlation could be detected
between the age of the female at first oviposition and
earliness or lateness in the season.
Records were also kept on the number of days that
elapsed between the various egg cases for 35 females.
‘As mentioned above, each fémale deposits during her
life-time from one to four cases; these, however, do not
follow in rapid succession, but at the following intervals:
Days Days Days
Roach between between between
Number Ist & 2nd 2nd & 3rd 3rd & 4th
1 6 15 —
2 12 ae Se
3 8 oe as
4 11 ae ou
The Biology of the Roach 65
Days Days Days
Roach between between between
Number Ist & 2nd 2nd & 3rd 3rd & 4th
5 10 ie ase
6 9 fe is
| 11 RG, aoe
8 6 15 st
9 19 be in
10 6 12 ya
11 11 ie ae.
12 8 16 og
13 8 sds ie
14 8 8 pee
15 8 we ee
16 10 13 ae
17 10 13 ae
18 9 12 mit
19 23 ae --
20 11 ies a
21 13 8 8
22 13 6 10
23 12 6 11
24 12 5 11
25 6 10 ==
26 4 7 -<
27 4 6 --
28 4 9 aia
29 12 -< a
30 11 =< mer
31 9 -= oe:
32 9 -- “
33 9 10 14
34 10 14 i
66 Trans. Acad. Sct. of St. Louts
These figures indicate that there is no appreciable dif-
ference in the time required for the formation of the
second, third or fourth egg case, and also that this inter-
val is practically the same (except for a few cases) as
the time elapsing between the last moult and the first
oviposition, as shown on page 63. Both sexes were often
kept together in the jars, and it is not known whether
mating occurred more than once. The eggs were all
fertile, but I suspect that these long lapses between egg
cases are probably due to the maturation of the eggs,
rather than a necessity of mating for each egg case.
An interesting observation from the biological stand-
point would be to test the fertility of the eggs in rela-
tion to one or more matings.
Period of incubation—The period during which egg
cases were deposited extended from the latter part of
May to the end of July. The period of incubation seems
unusually long. At the time of the emergence of the
nymphs, all leave the egg case at one time, instead of the
alternative of the emergence covering a number of days
for each case. The following data on the period of in-
cubation were collected on egg cases deposited from May
21st to June 11th:
Number of Number of
Days egg cases | Days egg cases
45 8 52 6
46 7 53 6
47 3 34 1
48 1 59 5
49 1 56 4
50 3 61 1
_ 8 Total___-54
The Biology of the Roach 67
Thus we see all of these 54 egg cases (exclusive of the
last, which might safely be regarded as abnormal), hav-
ing a period of incubation varying from 45 to 56 days.
The egg cases, as each was deposited, were placed in
Jelly glasses and kept in the third floor laboratory. It
seems to me that, if temperature has anything to do with
hatching, this should have accelerated their development,
for it was much warmer in the attic in July than in the
cellar where the eggs normally are laid.
Emergence of the young from the egg case-—When
first I witnessed the hatching of the roaches, I was much
surprised to find that their emergence is very similar to
that of the mantis, Stagmomantis carolina. The roach
egg cases are the well-known purses, the upper edge of
which shows a ridge or seam. By one of those miracles
which make marvelous every type of birth, this purse or
capsule is pressed open at this seam at the moment of
the emergence of the young, and the inmates are all
released at the same time, after which, the pressure
relieved, the capsule again closes and appears outwardly
exactly the same as before. If one examines a number of
egg cases previous to hatching time, and presses the ends
together with the fingers in an attempt to open this seam,
he will find them so tightly sealed that the case will mash
under the pressure before this ridge will open. At hatch-
ing time, however, they will yield readily, and at the
slightest pressure will open beautifully, revealing the
two halves of the case, each with the row of about eight
mummy-like organisms, perfectly white except the eye-
spots, which are black, and all arranged in two precise
rows, heads up. In about three minutes after opening
the case, all will have struggled out, and they are at once
extremely active. This reveals the fact that when the
egg case is deposited it has no opening, and only at the
68 Trans. Acad. Sci. of St. Louis
time of emergence does the opening gradually form.
One often wonders, upon picking up an empty egg case,
how it is possible for the sixteen young to have emerged
without leaving a break in the case. The mechanics is
extremely interesting. As in the Carolina mantis, all the
eggs are deposited with the head nearest the opening.
The eggs, being produced practically at one time, or at
least within forty-eight hours, all develop alike. In this
mummy stage preceding emergence, the caudal end of
the body tapers, while the greatest development is about
the head. Now with all of the enlargement occurring at
the fore part of the body, which is uppermost, it is easy
to see how sixteen enlarging fore parts would mechani-
eally and gradually open the exit, and simultaneously all
would slide out en masse and automatically the closure
would come together again.
Some writers think that the young secrete a liquid
which dissolves the cement in the seam of the capsule.
This process is not merely an emergence, but also a
real hatching, for in the process each insect issues from
its individual egg shell and also from the dotheca enclos-
ing a number of eggs. One often finds the thin, white
caps that cover the forepart of the insect caught in the
seam of the empty case, where the emerging nymphs have
left them behind. In addition to this cap each mummy is
completely enveloped in a thin, brown, papery bag, the
egg shell. It is very pretty to see, when one splits the
egg case in two parts, the two rows of heads facing the
opening, wriggling and writhing out of this paper shell,
and to see the two rows of heads, face to face, gradually
make their appearance. I have said that at this stage the
insect is completely white save the black eye-spots, but I
have discovered with the aid of the microscope that the
three teeth of each mandible are brown and seem to be
The Biology of the Roach 69
thoroughly chitinized, the only heavily chitinized part
- about the insect.
When this mummy is removed from this bag, the head
is inflexed, the legs, antennae and mouth-parts pressed
flat against the body, and the segments very distinct—in
fact, the whole organism looks like head and segments,
and not at all like the roach it will be in five minutes.
Even at this stage the two cerci are very prominent.
The roach works itself out of the egg shell by a series
of contortions, comes out through the opening and there
sheds a white membranous cap, which, no doubt, is for
the protection of the head. This frees its legs, and then
the newly born roach clumsily walks away. This behavior
is very much like that described for the praying-mantis.”
Another irregular piece of semi-chitinous covering
hangs from the tip of the abdomen, and this, I think, is
the shed skin that bound the legs and antennae close to
the body. With a few hard jerks this is left behind and
the roach, for only now does it really appear as a roach,
scampers briskly away. If the egg case is not artificially
opened, it indeed seems like magic to see almost simul-
taneously all heads protrude through the opening,
struggle for an instant to get the limbs free, kick the hind
legs loose from the adhering skin, and again simul-
taneously scamper away in all directions, all in less time
than it takes to tell.
In opening many egg cases I found all of them fertile
and in the same stage of development. In one case, how-
ever, amid healthy looking organisms, were two eggs
opposite each other, hard and brown, dead or infertile.
Number of young in each egg case.—Notes were made
on the number of nymphs that hatched from each of 43
egg cases. This number varied from 12 to 18, as follows:
*Trans. Acad. Sci. St. Louis, 22:1-57. 1913.
70 Trans. Acad. Sci. of St. Louts
No. in each Number of No. in each Number of
egg case egg cases egg case egg cases
12 2 16 15
13 5 1% 3
14 6 18 oe
19 10 Total____43
It is at once apparent that the mean number at which
nature aims is 16, and her deviations from that are not
significant; even the mortality in embryo is surprisingly
small.
Food of nymphs and adults—During the period of
observation an attempt was made to keep a record of
food materials which they accepted. Soon, however, the
list became so voluminous that it was apparent that it
would be more brief and simple to list the things which
they did not eat. This dwindled almost to nothing. They
showed a preference for the softer portions of the food
before them, and left the hard or chitinized portions, but
otherwise they ate everything of a vegetable or animal
nature, either fresh or decaying, which came in their
way.
Cockroaches are undoubtedly attracted to their food
by the sense of smell located in the antennae. Lloyd
Morgan * tells us that if their antennae be extirpated or
coated with paraffin, the roach no longer rushes to food,
and takes little notice of and will sometimes even walk
over blotting-paper moistened with turpentine or benzo-
line, which a normal insect cannot approach without
agitation.
Enemies.—Dermestes larvae often infest the egg cases
of the roach. Some large frogs of the edible variety
SAnimal Life and Intelligence, p. 257. 1891.
The Biology of the Roach 71
escaped from a tank in the cellar one night; when they
were recaptured and dressed the next day, the contents
of the stomachs revealed an astonishing number of
roaches in the large nymph stage. In nature the habitats
of the frogs and roaches are so remote that this relation-
ship can hardly be expected to exist. Some common gray
lizards kept in the laboratory also readily accepted the
roaches as food. ‘A neighbor told me that her chickens
greedily ate the roaches which she had caught in traps.
Woods’ says that the cockroach is a favorite food of
many animals and the hedgehog is so partial to it that
one of these animals is sometimes kept in the kitchen for
the express purpose of destroying these pests. A young
friend who is a nature student informs me that he fed
roaches of this species to a pet garter snake.
Roaches seem to require a larger supply of water than
do most species of insects, and they drink it with an evi-
dent enjoyment that is pleasant to see. Their food often
becomes mouldy, and fungus growths of various colors,
black, yellow, green or gray, cover it, but this seems not
to affect them in any way, and no increase in the mor-
tality could be noticed at these times. Very often, too,
their food, especially the starchy materials, becomes
covered with myriads of mites, Tyroglyphus lintneri
Osborn (identified by Mr. H. E. Ewing), but even these
do not affect the health or the mortality of the roaches in
the jars.
Method of concealing the egg cases.—The color of the
egg case as it leaves the abdomen is almost white. Some-
times the exposed part of the case becomes dark while
it is being carried, so a newly deposited case may have
every shade from white through a reddish pink to a red-
‘Insects at home, p. 237. 1872.
72 Trans. Acad. Sci. of St. Louis
dish brown. Eventually all change entirely to a reddish
brown.
Although the adults and nymphs of the roach may
appear in abundance about the house, the egg cases are
rarely seen. The reason for this was discovered in the
habits of those kept for close observation in the Jabora-
tory. In the great majority of cases, the mother very
deftly conceals the egg case, either by depositing it in a
erevice or by carrying earth and debris, sometimes for
considerable distances, and with this and her saliva,
daubing the new case until it is quite hidden from view.
This is so stealthily done that I have never been able to
eatch her in the very act, but the result leaves no doubt
that the act is deliberately and purposely done. Of
course the insects in their native habitat have every
chance to conceal the egg cases beyond all chance of our
ever finding them; even in glass jars, with a limited
amount of material with which to work, they succeeded
so well that in a few cases I myself did not discover
them until I had injured them in handling.
A large number of females carrying egg cases were at
different times placed in the jars, together with a small
amount of earth and rubbish, just to see what would
be their course of action. Out of 90 egg cases thus de-
posited, 36 were deposited in some place which afforded
some concealment, as crevices in bark, the cells of old
wasp nests, under a bit of paper or, in a few cases, in
slight depressions which, I was sure, the mother roach
had made in bark or clay, expressly to fit and receive her
dotheca. Many, if not the majority, of these were also
daubed. Of those which were dropped on the bottom of
the jars (this open space formed by far the greater part
of the available space), 38 were well hidden by having
been daubed with dirt and rubbish stuck on with saliva,
The Biology of the Roach 73
and only 16 were left uncovered. It is only reasonable
to surmise that the unnatural conditions of their life at
this time and my frequent interruptions may have hin-
dered the normal functioning of some of the mothers in
so delicate a task as this, and while the mothers in con-
finement concealed only 82 per cent of the egg cases,
in their own environment and undisturbed, they would
show a greater per cent of efficiency. As suggested
above, if the egg cases had been dropped by mere chance,
by far the greater part of them would have been on the
floor of the jars. In many cases they were found
crammed into crevices that must have been really dif-
ficult of access, and in more than one instance, three or
four lay end to end in the same cranny in the bark,
when nooks were scarce. In some cases we know posi-
tively that the mud must have been carried at least three
inches.
I repeat the assertion that the act of plastering the
egg cases must be deliberately done. While these
mothers were never caught in the act of this work, we
have the interesting details of this behavior of an allied
Species, Periplaneta americana Linn., by V. R. Haber.”
He says that at 2:50 a. m., the female began to scar and
roughen the surface of a cardboard in the cage. She
chewed and munched at the upper surface of the paste-
board until she had made quite an appreciable dent, not
dropping the pasteboard on the bottom of the cage, but
mixing the bits with a secretion from the mouth until all
became a damp mass. At 3:30, she crawled forward over
the scar with her abdomen bent anteriorly and ventral-
ward, probing about with the protruding dotheca until
she located the sear which she had just made. After sev-_
eral unsuccessful attempts at placing the egg case in the
10Ent. News 31:190-193. 1920.
74 Trans. Acad. Sct. of St. Louis
scar, because each time it rolled off, it finally fell and
bounced to the far side of the cage, whereupon she
promptly ran down and located it on the floor. She
cleaned it with her mouth-parts, coated its exposed sides
and ends with a secretion from her mouth and then
picked up loose bits of trash, with which she covered it.
She even went so far as to try to cover it with a piece of
paper. Girault” also found this habit of covering the
egg case with mud or bits of wood prevalent for Peri-
planeta australasiae Fab.
The mating habits of the roach.—In the proper sea-
son, a sudden switching on of the light often reveals a
pair of roaches in copulo; they are motionless, and are
united with their heads in opposite directions. The
mated pairs are found on vertical walls, as well as on
horizontal surfaces, but always in the same position.
While I have seen many mated pairs, and have seen some
attempts at mating, it still remains a mystery to me how
this position is accomplished; it must be through some
intricate movements. Mating always or nearly always
occurs in darkness, and it is most difficult to observe the
manoeuvers of these shy creatures, although by timing
the experiments and by switching on and off the lights,
one can get good records of the wooing. The ardor of
the courting male is not cooled when the lights are turned
on, but the modest little mate usually walks away and
breaks up the party.
In the preliminary movements the male gets directly
in front of the female, but turns so his head is away from
her (so that the two are in single file), with the rear tip
of his body directly in front of her face. He then walks
backwards, pushing his body underneath hers and at the
same time lifts his wings and holds them vertically,
NEnt. News 26:54. 1915.
The Biology of the Roach 75
thereby exposing the dorsal segments of his abdomen.
Sometimes a feeble attempt is made to flutter these ver-
tical wings in a way that suggests coquettishness. When
he has partly inserted his body under her, she slowly
walks on top of his back, touching or feeling the seg-
ments of his abdomen with her jaws and palpi. The cerci
of the male are protruding and his clasping organs ex-
tended and opening and closing like a pair of tongs, in
his attempt to grasp or feel segment after segment as
she crawls forward over him or as he crawls backward
under her (sometimes it is difficult to say just which is
really happening). These movements continue, and just
when one expects a culmination, the female breaks away
or the male makes a false step and the pair separate.
There must be some very interesting contortions occur-
ring before he makes the connections and faces about
completely from backing under the female to facing in
the opposite direction, and it is probable that many
attempts are made before this intricate task is accom-
plished. I hope some day, with better technique in the
way of light regulation and mirrors, to be able to get the
details.
There is actual courtship on the part of the males, and
there is actual rivalry between them, as will be seen from
the following experiment. On June 2, at 11:35 p. m, I
placed 9 chaste males about two weeks old (since they
had become adult) with a virgin female. Five minutes
later, upon switching on the lights, I found the female
had crouched under a bit of bark, and at the end of ten
minutes she had already mated with one of the males,
and several of the others, walking high up in the air
with legs outstretched like stilts (an extremely unusual
position which might be a way of trying to attract the
attention of the female), were pestering the pair. One
76 Trans. Acad. Sci. of St. Louis
male in particular seemed to be biting the pair apart, and
when I examined the spot after he had left I found that
he had bitten away a large portion of the wing of his
rival. I examined them at intervals for an hour, and
found the pair still in copulo; the next morning they
were separated, but there must have been some rivalry
between the males, for two others had the wings badly
bitten away. In another cage containing one female and
four males, similar rivalry occurred; the female was pas-
sive while the males ‘‘walked on stilts’? and protruded
claspers. After some little display, three of these males
retired and left her to the other suitor, which pursued
her for fifteen minutes before he attempted to back under
her.
C. L. Turner” says the process of copulation in the
cockroach is so rapid that the details cannot be followed;
it can only be said that the male shoves his body under
that of the female, and accomplishes the transfer of sper-
matozoa in a few seconds. All those which I observed
were much more deliberate, and in certain instances sev-
eral hours were spent in actual mating.
I have recorded* very similar courtship behavior for
the wild roach, Parcoblatia pennsylvanica DeGeer, but
here in the only case observed the female assumed the
aggressive role.
During the night males often go through mating ma-
noeuvers with other males.
Ezxudation.—In working with adults one soon notices
at times a gathering of a thick, milky jelly on the ventral
surface that seems to ooze out at the lines of demarca-
tion of the last two segments. This is an almost trans-
parent, slime-like substance of sufficient gumminess to
22Ann. Ent. Soc. Amer., 9:122. 1916.
18Trans. Acad. Sci. St. Louis., 24:57. 1922.
The Biology of the Roach 77
hold itself together for many days, and is very notice-
able on the black backs of the roaches. Just exactly
where it comes from or what its function is, I do not
know, but it probably oozes out from pores in thin chitin
at the joints of segments and probably functions in some
way in the mating or reproductive processes. This ma-
terial was never seen to gather on the bodies of the males
or on the mated females, but only on the virgin females.
It is not wholly accurate to say that the mated females
never showed this secretion; I should say, rather, that
this material did not appear on the bodies of hundreds
of females caged together with males, but at the same
time in a group of virgin females that were segregated,
almost all of them had varying amounts of this slime
that exuded from the dorsal folds of the last few
segments.
Miscellaneous activities—Roaches are negatively
phototactic and positively thigmotactic. Writers gen-
erally agree that roaches are gregarious, but it seems
to me that their gregariousness has more or less been
thrust upon them; being negatively phototactic and posi-
tively thigmotactic, and with large numbers of them seek-
ing the same environment in limited space and hiding
places, one can see how easy it is for the roach to appear
as gregarious, but there probably is no more social ten.
dency in the life of the roach than in any other member
of the Orthopterous group.
While the roaches are dark-loving creatures, there is
a behavior that indicates a response which is so aptly
called differential sensibility. This can be tested by sud-
denly lighting up a roach infested cellar. The creatures
run frantically until they come within the shadow of a
post; there they stop short, and may easily be picked up.
They seem certain of safety when in this darkness,
78 Trans. Acad. Sct. of St. Lows
although what they respond to is not real darkness, but
only a less light condition. In the same way, I have seen
roaches escape into crevices; when their bodies touched
the crack they were satisfied, even though they were ex-
posed to full view in the light; their condition of positive
thigmotropism satisfied their feelings in the matter of
protection from danger, and they did not venture deep
into the crevice. Here, however, it seems that light and
sight were factors that prevented me from picking them
up, for when attempts were made they crawled deeper
into the crevices, whereas in the dark shadow on the floor
an attempt was seldom made to escape because the roach
could not see my approach.
Roaches are very sensitive to odors or air conditions.
One may handle the jar without disturbing the inmates
if the jar is covered, or one may even remove the glass
cover and come very near to look in if the breath is
deflected, but to breathe, even gently, down into a jar
at once creates a panic among the roaches. This simple
test shows them to be susceptible to odor or air vibra-
tions more than to sight.
Their coming out of hiding does not seem to be regu-
lated entirely by the quietness of the place. On many
occasions I have found, when a house had been closed
for several hours, roaches were out to feed as early as
8:30 or 9 o’clock; on other occasions they may be found
out foraging when the room has been darkened only a
few minutes, when the noises of the city have by no
means subsided. On the other hand, during the night
when everything is quiet and one expects them to be out
in abundance, one may find few or none. It seems they
feed as early in the evening as possible and then retire.
The Biology of the Roach 79
I have never seen male roaches use their wings for
flying. I have thrown them up in the air; they seem to
have no control over the fall; they did not even open the
wings to act as a parachute, but turned many somer-
saults in the air and hit the earth with a thump. The
males can, however, make their escape by backward pro-
gression as rapidly as by traveling forwards.
%
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Transactions of the ease - Science of St. Louis
ae ntitgy
Volume XXV, No. 5 we
NOTES ON THE STENOMORPHIC FORM
OF THE SHIPWORM
W. F. CLAPP
| vy eS Gur BORg,
Issued January, 1925 eS e Wy
. [ RB ‘
NOTES ON THE STENOMORPHIC FORM OF
THE SHIPWORM.
W. F. Crapp.
The term stenomorph has been given by Dr. Bartsch
(Bartsch, 1923, p. 330) to those specimens of shipworms
which are abnormal in being dwarfed, although they may
be sexually mature. Dr. Bartsch discovered the fact
that this form is frequently found in small pieces of wood
and inferred that it is caused by the cramped habitat
of the individual. A study of the stenomorphic specimens
found in the test-blocks placed by the Committee on
Marine Piling Investigations of the National Research
Council (Clapp, 1923, p. 31) has added further evidence
of the conditions which cause this peculiar form.
The shell of the stenomorphic specimen (pl. 4, fig. 1)
may generally be distinguished from the normal speci-
men of the same species by its smaller size and greater
shell thickness, as well as by the proportionately greater
number of denticulate ridges on both the anterior and
the anterior-median areas of the shell. The internal
blade or apophysis is greatly broadened and strength-
ened (pl. 4, fig. 2). There is, however, no sharp
dividing line between the stenomorph and the normal
Specimen of the same species. Many large specimens,
with tubes 250 millimeters or more in length, show
traces of stenomorph characters in those portions of
the shell last secreted. There is a well-graded series
between these large, nearly normal specimens and those
typical stenomorphs of the same age which possess tubes
only ten millimeters long. In the typical stenomorph
the auricle is entirely lost, being overlaid by the an-
82 Trans, Acad. Sci. of St. Louis
terior-median area, since the denticulate ridges of this
area completely cover the entire median area and also
the auricle to its posterior edge. The anterior area
is also proportionately large, the denticulate ridges of
that area being added to its ventral edge until it is
nearly as large as the anterior-median area both in
length and in height. In a young normal specimen of
Teredo navalis, the ridges of the anterior area are
approximately one-fourth the width of the intervening
spaces (pl, 4, fig. 3). In a typical stenomorphic speci-
men of the same species and of the same size, the den-
ticulate ridges of the anterior area are so closely
crowded that the spaces between the ridges are reduced
to mere threads (pl. 4, fig. 4). This dwarfed form, while
the same age as a large normal individual, may be but
one-tenth of its size. The embryos in the gills of a
sexually mature stenomorphic specimen are much less
numerous than are the embryos in a normal specimen
of the same species, but in all other respects the embryos
are identical.
At many of the places where test-blocks have been
placed by the Committee on Marine Piling Investiga-
tions, these stenomorphic specimens are frequently
found in the wood, and since the blocks used in 1922
were small (2 inches by 4 inches), it was thought that
the abnormality was caused by the cramped quarters.
As Dr. Bartsch states (loc. cit.), they are frequently
found in laths and other small pieces of wood, but that
some factor other than the size of the wood enters into
the result, is shown by the fact that laths and small
pieces of wood frequently contain large and perfectly
normal specimens. For example, Teredo navalis causes
considerable damage to the wooden lobster pots in the
Notes on the Stenomorphic Form of the Shipworm 83
waters south of Cape Cod. The laths of which the
lobster pots are made are often completely riddled in a
few weeks. The number of stenomorphic specimens in
these laths is no greater proportionately than one finds
in heavy piling and the average size of a larger number
of specimens removed from the laths is the same as
that of specimens removed from big timber. ‘Also, large
blocks, made up of many shingles bolted more or less
tightly together (Mark, 1924, p. 266, footnote 3, p. 268,
fig. EZ), were placed at fifty or more stations from Massa-
chusetts to Texas. In some of these shingle blocks,
stenomorphic specimens occurred, but in the majority
the specimens were not stenomorphic, but nearly or en-
tirely normal. Because of the fact that the average
shipworm will endeavor to avoid crossing a large crack
or open area in the wood, as long as progress is possible
in some other direction, many of the shipworms in the
shingle blocks made more or less of an effort to remain
within a single shingle, the effort depending entirely
upon how firmly the shingles were bolted together.
When the specimens entered the thicker end of the
shingle and advanced with the grain of the wood toward
the thinner end, this effort to refrain from crossing to
the adjacent shingle, resulted in a peculiarly abnormal
form, in which the tubes are excessively attenuate.
Many of these specimens were found which had attained
a tube length of 400 millimeters in three months, but,
owing to the thinness of the wood, had increased the
diameter of the tube very slowly. The anterior ends of
many of these long tubes were frequently but five milli-
meters and occasionally only four millimeters in
diameter. The shipworm is able to contract its body
into one-half of its normal length. The body is firmly
attached to the wall of the burrow only at the extreme
84 Trans. Acad. Sci. of St. Louis
posterior end. In an effort to locate thicker wood an
individual may occasionally withdraw its shell and the
anterior portion of its body from the anterior portion
of the tube, forming a new tube in a different direction.
At the point of separation from the original tube a
diaphragm is formed completely sealing the abandoned
tube. This operation may be repeated several times by
a single individual (pl. 5, fig. 3). Tubes of normal speci-
mens of the same species which are 400 millimeters in
length are approximately twelve millimeters in diameter
at the anterior end. This abnormal form which occurs
in thin wood, however, possesses none of the characters
of the stenomorph. The thickness of the shell, arrange-
ment of the denticulate ridges and the proportionate
sizes of the anterior and anterior-median areas, being
the same as in normal specimens, the abnormality
caused by the thickness of the wood can be seen only in
the excessively narrow form of the tube. The large
proportion of abnormal specimens with attenuate tubes
and the small number of true stenomorphs found in the
shingles, would tend to show that the stenomorph
characters are not caused by the size of the wood.
Also, many of the small test-blocks used in 1922 con-
tained in one portion of the block many hundreds of
stenomorphic specimens, while in another equally large
portion of the block there were only relatively a few
normal specimens of the same species. For example,
small test-blocks from Galveston, Texas, contained for
several weeks during the summer of 1922, in one-half
of the block, hundreds of specimens of stenomorphic
Bankia gouldi, few being more than twenty millimeters
in length, whereas in the other half of the block there
were always two, three, or more large normal 200-300
Notes on the Stenomorphie Form of the Shipworm 85
millimeter specimens of the same species. Furthermore,
at the same locality the increase in the size of the test-
blocks in 1923 to 4x4x6 inches did not change the result
in the slightest degree, the proportion of stenomorphs
being as great and the average size of the stenomorphic
specimens being the same in the large blocks as in the
small. Also, several of the numerous specimens of piling
received from various localities on the Atlantic Coast,
some of which were 18 inches or more in diameter,
showed the same stenomorph form, not separable from
those found in the small test-blocks.
From the above facts it is evident that the size of
the wood is not necessarily the deciding factor in the
production of the stenomorphs, for if it were, steno-
morphs would not occur in large piles where plenty of
wood remained unoccupied. In a great majority of the
pieces of wood containing stenomorphic specimens which
I have examined, I have found that the conditions at the
time of the attachment to the wood of the embryo, were
apparently very favorable. Many embryos entered the
wood at the same time, frequently thirty or forty to
the square centimeter. A large majority survived, but
many specimens, owing to the fact that some of their
neighbors entered the wood slightly earlier, or were
more active in boring than they, soon found themselves
unable to advance in any direction, for the reason that
they were completely surrounded by the tubes of other
specimens. The shipworm will never break through the
partition of wood, however thin it may be, which sepa-
rates it from a neighboring tube, whether the occupant
of that tube be living or dead. These confined specimens
continued to live for as long a period as those more
fortunate individuals which were not checked in their
86 Trans, Acad. Sci. of St. Louis
growth. Since it was not possible for them to increase
the length or diameter of the tube or the size of the shell,
the only noticeable activity was in the regular addition
of denticulate ridges and an increase in the thickness
of the shell, this peculiar growth rendering them almost
unrecognizable. In view of the fact that the shipworm
is supposed to be able to obtain more or less nutriment
from the wood into which it bores (Dore and Miller,
1923), (Harrington, 1921), it is interesting to note that
the stenomorphic specimen, at least, is not dependent
upon the wood for food, for living specimens of steno-
morphic Teredo navalis have been found with a total
tube length of only ten millimeters and yet with a known
age of ten months. A normal Teredo navalis during the
active season, will form a ten-millimeter tube in less
than two weeks time. That the animal of the steno-
morph should continue to produce denticulate ridges
when there is no more available wood upon which the
denticles can be used, is even more interesting.
I wish to express my appreciation for the assistance
which has been given to me in my study of the shipworms
at the Massachusetts Institute of Technology, through
the kindness of Professor Samuel C. Presscott. Ex-
cellent laboratory facilities, photographic apparatus,
etc., have been freely placed at my disposal.
I am also indebted to Mr. Nelson M. Fuller for the
excellent photographs and for other assistance in my
work.
TrANS. Acap. Sci. oF St. Lours, Voit. XXV Pirate IV
1G 2
Notes on the Stenomorphic Form of the Shipworm
EXPLANATION OF PLATE
Piate IV
External view of both valves of a ee specimen
Teredo navalis. Actual size .94 mm
Internal view of both valves of a stenomorph specimen
Teredo navalis. Actual size .94 m
External view of h valves of a normal specimen
: .3 mm.
ew of bot
Teredo navalis. Actual size 1
External view of left valve of a stenomorph specimen
Bankia gouldi. Actual size 2.7 m
87
of
of
of
Pallets of stenomorph Teredo navalis shown in fig. 1. Actual
i 5 mm.
88 Trans, Acad. Sci. of St. Louis
EXPLANATION OF PLATE
PLaTe V
Fig. 4 secon Scene many stenomorphic specimens of Teredo
lis from Warren, R. I.
Fig. 2. aa showing attack of normal Teredo navalis from Fall
River, Mas
Fig. 3. Tube of a ee gouldi in a shingle block.
TRANS. Acab. Sci. oF St. Louis, Vot. XXV PLATE V
Notes on the Stenomorphic Form of the Shipworm 89
BIBLIOGRAPHY
1921. Harrington, C. R.—A note on the physiology of the shipworm
Teredo norvegica). The Biochemical Journ. Vol. XV. No. 6,
741, fig. 1.
pp. 736-
1923. Bartsch, Sg aes ali a new term in taxonomy. Science
N. S. Vol. 57, p. 330.
1923. bea tok —— Pere ew species of Teredo from Florida. Proc.
Bos <. Net. Hist. Vol. 37, No. 2, pp. 31-38, pl. 3-4.
1923. Dore, W. H. and R. C. Miller—The digestion of wood by Teredo
navalis. Univ. of Cal. Publ. Zool. Vol. 22, pp. 383-400, pl. 18.
1924. Mark, E. L.—Marine borers in Bermuda. Proc. anes Acad. Arts
and Sci. Vol. 59, No. 11.
Coy Rm
: at
Transactions of the Academy of Science of St. Louis
Volume XXV, No. 6
SUPPLEMENT TO THE CATALOGUE OF
ARKANSAS PLANTS
J. T. BUCHHOLZ and E. J. PALMER
Se
<9
ae
< »
Issued June, 1926 19.19%) Py
SUPPLEMENT TO THE CATALOGUE
OF ARKANSAS PLANTS.*
J.T. Bucunouz ano EB. J. Paumen.
More than thirty-six years have elapsed since the pub-
heation of a Catalogue of the Flora of Arkansas. We
are presenting here the first supplement to the List of
Arkansas Plants by Branner and Coville (35). Many
plant names have been changed since this publication, but
we are not attempting to include names in this list where
mere changes in synonymy are involved, except in a few
instances where these changes are such as would not be
included with the synonymy given in current botanical
manuals. Our supplementary list of plants presented
here consists almost exclusively of additions to the for-
mer catalogue and includes also the species reported by
various other botanists, both in publications appearing
since 1890, and from reports rendered privately to the
Department of Botany of the University of Arkansas
within the last six years.
One or more collection numbers are given where pos-
sible. However, collections made by J. T. Buchholz were
usually not numbered, and when thus marked, all num-
bers used by this observer are below 1000. Nearly all
of these specimens are in the Herbarium of the Uni-
versity of ‘Arkansas. Collection numbers of Arkansas
plants by E. J. Palmer are those given with numbers
above 4000, unless otherwise stated, and duplicate speci-
mens of many of these plants are in the Herbarium at
Fayetteville. Most of the early collections by Palmer
*Research Paper No. 22, Journal Series, University of Arkansas.
(91)
92 Trans. Acad. Sci. of St. Lows
are to be found in the Herbarium of the Missouri Botan-
ical Garden, while a complete set of collections of all
ligneous species with Palmer numbers is to be found in
the Herbarium of the Arnold Arboretum.
The flora of Arkansas has been largely neglected by
botanists during the last thirty-six years. Those who
have collected within the State have usually been re-
warded by the discovery of many new and rare species
of plants. It is safe to predict that as this flora becomes
better known very many more additions will be made,
and with a more careful and critical study of the material
many new species and varieties will be added.
The Catalogue of Plants by Branner and Coville (34)
included 1610 ferns and seed plants. A small reduction
must be made from this list for plants erroneously re-
ported or those now represented under new names among
our 534 additions. However, it may safely be estimated
that the known ferns and seed plants of Arkansas now
comprise about 2100 species.
The earliest naturalists to visit and report on the
vegetation of the area included in this state, of which
there are available records, were Bradbury (1) in
1809-11, Schooleraft (2) in 1818-1819 and Nuttall in 1819.
All of these early explorers gave valuable and interest-
ing accounts of the geography and vegetation, but the
celebrated botanist Nuttall described the vegetation in
greatest detail and published valuable contributions in-
cluding the descriptions of many new species of plants
(4, 5, 7). Nuttall’s Journal of Travels Into ‘Arkansa
Territory (3) is considered a classic in the literature of
travel, and this together with his Collection Towards a
Flora of the Territory of Arkansas (7), are noteworthy
Supplement to Catalogue of Arkansas Plants 93
contributions to the flora of the West. His Flora of
Arkansas was never completed, but he mentioned many
plants in his Journal of Travels, and described others in
his Flora and other contributions (4, 5). The works of
Bradbury, Schoolcraft and Nuttall constitute the earliest
reliable records on the flora of Arkansas.
In 1834 Featherstonehaugh traveled through Missouri
and Arkansas from the Missouri to the Red River.
Though his report (6) was largely geological, he made
frequent mention of many species of plants observed and
the types of vegetation encountered.
In 1859 Professor Leo Lesquereux made a study of the
fossil and recent flora of the state for the Second State
Geological Survey, and gave a description of the botani-
cal features of the northern and northwestern counties
and a catalogue of Arkansas plants (8), including those
observed and reported by Nuttall. As Lesquereux did his
work during October-December and covered very exten-
Sive territory, he probably did not assemble an herba-
rium.
To this catalogue Butler added a list (9) of over a
hundred species in 1877. Prof. F. L. Harvey published
many short articles snd notes in the Botanical Gazette
(10-15, and 17-29) between 1880 and 1885, including a
more comprehensive treatise on the Ferns of Arkansas
(17-18) and one on the arboreal flora (32). Harvey
began to assemble an herbarium. A large portion of his
collection is still preserved at the University of Ar-
kansas, and many of his duplicates were exchanged with
other herbaria.
Professor R. Ellsworth Call, who was engaged in a
Study of the geology of Crowley’s Ridge, prepared two
94 Trans. Acad. Sci. of St. Louts
special reports on the native forest trees of this region
(33, 37). He probably did not assemble any collections.
J. Francis Williams made a special study of the timber
of the Magnet Cove region (38) and reported a definite
relation between the vegetation and the underlying
igneous rocks, confirming a feature previously mentioned
by Featherstonehaugh, but apparently not observed by
Harvey (21).
In the Annual Report of the Arkansas Geological Sur-
vey for 1888 Branner and Coville (35) published the
most recent comprehensive catalogue of Arkansas plants,
a publication which appeared in 1891. This also in-
cluded a discussion of the general botanical features of
the state (35). Coville made some collections which are
on deposit in the U. S. National Museum, but the bulk of
the work done for the plant list seems to have been based
on field notes. The Harvey collection at Fayetteville
was worked over by Professor Simonds in preparing
the Branner and Coville catalogue.
An extensive collection made by Mr. E. N. Plank, who
lived at Decatur, Arkansas, for many years, was acquired
by the New York Botanical Garden. The Plank collec-
tion contained many Arkansas plants, but he seems not
to have published a list of these except in a single brief
paper: ‘‘Concerning the Plants of Southwestern Ar-
kansas’’ (41). The material collected by Plank and in-
cluded in this report has been consulted at the herbarium
of the New York Botanical Garden.
In an ecological study, Dr. S. M. Coulter, (43) re-
corded a number of swamp species new to Arkansas.
Similarly Dr. Roland M. Harper (47-49) has made
records of a number of species characteristic of our
Supplement to Catalogue of Arkansas Plants 95
prairie and the coastal plain region of southeastern ‘Ar-
kansas, containing names new to the state. More re-
cently Dr. Harper and Rev. H. E. Wheeler sent us speci-
mens representing unreported species, as credited to
them. Others contributing data, specimens, or assist-
ance in finding favorable collecting stations are: Mr.
Ralph Shreve of Farmington, Prof. D. Demaree of Hen-
drix College, Conway; Mr. W. W. Ashe of the Forest
Service; staff members of the Agricultural Experiment
Station, Fayetteville; some of the county agricultural
agents and a number of former students of the Univer-
sity of Arkansas. These miscellaneous unpublished
records are included here with the name of the observer.
From these, from some of our own publications, and for
the most part from our other unpublished records, this
list of additions to the catalogue of Arkansas plants has
been prepared. A list of publications pertaining entirely
or in part to the flora of Arkansas is given below. This
is represented by a complete collection of these publica-
tions, nearly all bound in eight volumes deposited in the
University of Arkansas Library.
PTERIDOPHYTA.
Polypodiaceae:
Polypodiwm virginianum L. (formerly reported
as P. vulgare L.) Fayetteville, Washington Co.;
Jasper, Newton Co.; Magazine Mountain, Logan
Co., probably the southwestern limit of the species.
Pteridium aquilinum var. pseudocaudatum Clute.
Savoy, Washington Co.
Pellaea glabella Mett. Bentonville (near Bella
Vista), Benton Co. |
96 Trans. Acad. Sci. of St. Lowis
Asplenium platyneuron forma serratum (KE. 8.
Miller) R. Hoffman. On chert and limestone rocks,
with the species.
Polystichwm achrostichoides forma incisum (Gray)
Gilbert. With typical form, chert hills, Northwest
Arkansas, 7
Dryopteris spinulosa (Muell.) Kuntze. On north
slope, Magazine Mountain, Logan Co. (D. M.
Moore).
Woodsia scopiina D. C. Eaton. On sandstone
ledges, north side of Magazine Mountain, Logan Co.,
near the summit.
Dennstaedtia punctilobula (Mich.) Moore. On par-
tially shaded damp sandstone cliffs, north side of
Magazine Mountain, Logan Co. 24142.
Ophioglossaceae:
Ophioglossum Engelmanni Prantl. 5376 Fulton,
Hempstead Co., clay barrens; also in glades and
rocky barrens of the Ozark region.
Botrychium obliquum var. tenuifolium (Underw.)
Gilbert. 24008 McNab, Hempstead Co.
Botrychium obliquum var. dissectwm (Spreng-)
Clute. Farmington, Washington Co. Low woods,
rare, occurs with the species.
Equisetaceae:
Equisetum hyemale var. intermedium Fat. fide J-
H. Schaffner. On calcareous deposits near spring,
Jasper, Newton Co.
Supplement to Catalogue of Arkansas Plants 97
Equisetum praealtum Raf. (E. robustum (A. Br.)
A. 'A. Eaton.) Includes the plants formerly referred
to E. hyemale. Along sandy river banks.
Isoetaceae:
Isoetes Butleri Engelm. Little Rock, on ‘‘Pu-
laskite,’’ Fourche Mountain; Eureka Springs, Car-
roll Co. (Bush).
Isoetes melanopoda J. Gay. In springy ground,
top of Magazine Mountain, Logan Co.; 24921 Hot
Springs, Garland Co.; Farmington, Washington Co.
MONOCOTYLEDONAB.
Sparganiaceae:
Sparganium americanum Nutt. Prairie View, Wash-
ington Co.; 10598 Washington, Hempstead Co. ; 8358
Ashdown, Little River Co.
Alismaceae:
Sagittaria longirostra (Micheli) J. G. Sm. Fayette-
ville, Washington Co.; 10599 Washington, Hemp-
stead Co.
Sagittaria papillosa Buch. 25043 Hazen, Prairie Co.
Graminae:
Erianthus strictus Baldw, Fulton, Hempstead Co.
(4386 Greenman).
Paspalum distichum L. Fulton, Hempstead Co.
(4387 Greenman).
Paspalum circulare Nash. 6363 Beaver, Carroll Co.
98
Trans. Acad. Sci. of St. Louis
Paspalum elatum L. C. Rich. Corning, Clay Co.
Paspalum laeviglume Scribn. 26210 Cotter, Baxter
Co.
Paspalum Muhlenbergii Nash. 6093 Corning, Clay
Co.; 6370 Beaver, Carroll Co.; 8175 Fayetteville,
Washington Co.
Panicum Boscii Poir. 8426 Benton, Saline Co.
Panicum commutatum Schultes. 25015 Little Rock;
24880 Hot Springs, Garland Co.
Panicum Helleri Nash. Eureka Springs, Carroll Co.
Panicum huachucae Ashe. 24827 Magazine Moun-
tain, Logan Co.
Panicum linearifolium Seribn. 5611 Eureka Springs,
Carroll Co.; 25178 Shirley, Van Buren Co.
Panicum lucidum Ashe. 25169 Shirley, Van Buren
Oo.
Panicum mutabile Scribn. & Smith. 24872 Hot
Springs, Garland Co.
Panicum polyanthes Schultes. 25053 Hazen, Prairie
Co.
Panicum Scribnerianum Nash. 5558 Eureka Springs,
Carroll Co.
Panicum tennesseense Ashe. Westfork, Washing-
ton Co.
Panicum Werneri Scribn. 5610 Eureka Siena:
Carroll Co.; 25024 Hazen, Prairie Co.
Echinochloa colona (L.) Link. 26727 McNab,
Hempstead Co.
Supplement to Catalogue of Arkansas Plants | 99
Digitaria Ischaemum Schreb, (Digitaria humi-
fusa Rydb.) 4564 Eureka Springs, Carroll Co.
Digitaria villosa (Walt.) Ell. Arkadelphia, Clark
Co. (fide H. R. Rosen).
Axonopus furcatus (Fluegge) Hitche. 10594 Wash-
ington, Hempstead Co.
Setaria viridis (L.) Beauv.
Holcus halepensis L. (Sorgum halepense Pers.)
Naturalized in waste places and widely distributed.
Sporobolus canovirens Nash. 6908 Harrison, Boone
Co.
Sporobolus cryptandrus (Torr.) Gray. 6055 Corning,
Clay Co.
Sporobolus pilosus Vasey. 4374 Eureka Springs,
Carroll Co. |
Sporobolus vaginaeflorus Torr. Fide Standley
and Hitchcock, from specimens by County Agr.
Agent.
Agrostis Elliottiana Schultes. Central Arkansas,
from District Agr. Agent.
Eragrostis glomerata (Walt.) Dewey. El Do-
rado, Union Co.
Bromus secalinus L. Warren, Bradley Co.
Bromus tectorum L, Fayetteville, Washington Co.
Sphenopholis pallens (Spreng.) Seribn. 25168
Shirley, Van Buren Co.
Glyceria septentrionalis Hitche. 25046 Hazen,
Prairie Co,
Trans. Acad. Sci. of St. Louts
Elymus arkansanus Scribn. & Ball. North Ar-
kansas, District Agr. Agent.
Elymus australis Scribn. & Ball. Fayetteville,
Washington Co.; 8126 Benton, Saline Co.
Elymus glabriflorus (Vasey) Scribn. & Ball.
Lake Village, Chicot Co.
Cyperaceae:
Cyperus compressus L. 14641 McNab, Hempstead
Co.
Cyperus cylindricus (Ell.) Britton. Fulton, Hemp-
stead Co. (fide J. M. Greenman).
Cyperus ferax Rich. 26747 Fulton, Hempstead Co.
Cyperus Houghtonti Torr. Fayetteville, Washington
Co.
Cyperus hystricinus Fernald. 14657 McNab, Hemp-
stead Co.
Cyperus lancastriensis Porter. 8124 Benton, Saline
Co.
Cyperus pseudovegetus Steud. 6061, 8026 Fulton,
Hempstead Co.
Cyperus rivularis Kunth. 4565 Eureka Springs,
Carroll Co.
Cyperus speciosus var. ferruginescens (Boeckl.)
Britton. Fayetteville, Washington Co.
Eleocharis Torreyana Boeckl. Lonoke Co.. woods
of second bottoms.
Fimbristylis Baldwiniana Torr. Stuttgart, Ar-
kansas Co. (H. R. Rosen). A troublesome weed of
rice fields.
Supplement to Catalogue of Arkansas Plants 101
Fimbristylis castanea var. puberula (Michx.)
Britton. 5531 Eureka Springs, Carroll Co.; Hazen,
Prairie Co., and Stuttgart, Arkansas Co., where a
troublesome weed in rice fields.
Fuirena hispida Ell. 8099 Gifford, Hot Springs
Co.
Fuirena simplex Vahl. 6002 Cotter, Baxter Co.;
8370 Ashdown, Little River Co.
Scirpus debilis Pursh. Lonoke Co., Portland clays
(rice land soil).
Rynchospora corniculata (Lam.) Gray. 8070 Arka-
delphia, Clark Co.; 10520 Doddridge, Miller Co.
Rynchospora cymosa Ell, South Arkansas, District
Agr. Agent.
Rynchospora gracilenta Gray. 8095 Gifford, Hot
Springs Co.
Rynchospora macrostachya Torr. 6071 Corning,
Clay Co.
Rynchospora Plankii Britton. Benton Co. (type
locality) ; Washington Co.
Carex arkansana Bailey. Arkansas River Valley.
Carex Bicknellii Britton. 25060 Hazen, Prairie Co.
Carex cherokeensis Schwein. 27104a Conway, Faulk-
ner Co.; Pulaski Co., Roland M. Harper.
Carex communis, Bailey. 26930 Magazine Mountain
Logan Co.; 27098 Jasper, Newton Co.
Carex crus-corvi Schuttlw. Corning, Clay Co.
Carex debilis var. Rudgei Bailey. 27142 Conway,
Faulkner Co.; 24938 Lonsdale, Garland Co.
102
Trans. Acad. Sct. of St. Louis
Carex festucacea var. brevior (Dewey) Fernald.
27107a Conway, Faulkner Co.
Carex glaucodea Tuckerm. 25069 Hazen, Prairie
Co.; 24874 Hot Springs, Garland Co.
Carex gynandra (Schwein.) Boott. 24927a Lons-
dale, Garland Co.
Carex hormathodes var. Richiit Fernald. 27007
Tontitown, Washington Co.
Carex intumescens Rudge. 24990 Fulton, Hemp-
stead Co.
Carex Jamesit Schwein. Fayetteville, Washington
Co.; 5581 Beaver, Carroll Co.
Carex Joorii Bailey. 24004 Fulton, Hempstead Co.
Carex awe Michx. 24747 Fayetteville, Wash-
ington C
Carex laxiflora var. gracillima Boott. 24746 Fay-
etteville, Washington Co.; 24452 Hot Springs, Gar-
land Co.
Carex laxiflora var. latifolia Boott. 6038 Rush,
Marion Co.
Carex leptalea Wahlenb. 27132 Lofton, Hot Springs
Co
Carex louisianica Bailey. 27106 Conway, Faulk-
ner Co.; Lonoke Co., heavy clay soil in woods.
Carex lupulina var. pedunculata Bailey. 6047 Corn-
ing, Clay Co.
Carex Meadii Dewey. 5533 Eureka Springs, Carroll
Co.
Supplement to Catalogue of Arkansas Plants 103
Carex Muhlenbergii var. enervis Boott. 5552
Kureka Springs, Carroll Co.; 24815 Magazine Moun-
tain, Logan Co.
Carex oligocarpa Schkuhr. 27036 Goshen, Wash-
ington Co.; 24849 Magazine Mountain, Logan Co.
Carex platyphylla Carey. 26929 Magazine Moun-
tain, Logan Co.
Carex Sartwellii Dewey. Lonoke Co.
Carex scirpoidea Michx. 24928 Lonsdale, Gar-
land Co.
Carex echinata var. angustata (Carey) Bailey.
27105a Conway, Faulkner Co.
Carex stricta var. decora Bailey. 24745 Fayette-
ville, Washington Co.
Carex tetanica Schkuhr. 27041 Eureka Springs,
Garland Co.
Carex torta Boott. 26898 Lawrence, Garland Co.
Commelinaceae: |
Tradescantia brevicaulis Raf, Fayetteville, Wash-
ington Co. On sandstone, cap rock of high hills.
Tradescantia hirsuticaulis Small. Hot Springs,
Garland Co. (Roland M. Harper). Dry woods near
summit of West Mountain.
Tradescantia occidentalis (Britton) Smyth. 25067
Hazen, Prairie Co.; Lonoke, Lonoke Co.
Tradescantia reflexa Raf. England, Lonoke Co.
Commelina communis L. (C. nudiflora Gray).
Texarkana, Miller Co.
104 Trans. Acad. Sci. of St. Louis
Juncaceae:
Juncus aristulatus Michx. Hazen, Prairie Co.
(Roland M. Harper).
Juncus brachycarpus Engelm. 25065 Hazen.
Prairie Co.; Jonesboro, Craighead Co., and 8S. E.
Arkansas. (Roland M. Harper.)
Juncus Dudleyi Wiegand. 5995 Cotter, Marion Co.
Luzula campestris var. bulbosa A. Wood (Jun-
coides bulbosum (Wood) Small). Most of our
plants, probably all, belong to this variety or
species. N. W. and central Arkansas.
Liliaceae:
Allium stellatum Ker. West Fork, Washington
Co.; 4370 Eureka Springs; Sulphur Springs, Ben-
ton Co.
Iilium superbum L. Wet meadows, Fayetteville,
Washington Co. (C. Chandler).
Stenanthium robustum Watson. Fayetteville, Wash-
ington Co.; 8205 Brentwood, Washington Co.
Toxicoscordion Nuttallii (Gray) Rydb. 24929 Lons-
dale, Garland Co.; Magazine Mountain, Logan Co.;
Rudy, Crawford Co.
Trillium pusillum. Michx. Valley of Osage Creek
and Monte Ne region, Washington and Benton Co.;
War Eagle Creek Valley, Madison Co. Occurs in
cherty soil in woods (rare).
Trillium Underwoodii. Small. Near Batesville,
Independence Co. (‘‘Cherty slope near Polk
Bayou’’—Roland M. Harper).
Smilax hispida Mihl. 10558 Cotter, Marion Co.;
22207 Van Buren, Crawford Co.
Supplement to Catalogue of Arkansas Plants 105
Dioscoreaceae:
Dioscorea villosa var. glabra. Lloyd. 5945 Cot-
ter, Marion Co.
Amaryllidaceae:
Hypoxis hirsuta (L.) Coville (H. erecta L.) 25027
Hazen, Prairie Co. Also frequent in Ozark region.
Iridaceae:
Iris cristata Ait. (Formerly reported as I. verna
L.) Cherty hills in woods north and west of Fay-
etteville, Washington and Benton Co.; similar situa-
tions Madison and Carroll Cos.; top of Magazine
Mountain, Logan Co. A form with white flowers has
been collected at Farmington and Hot Springs.
Iris foliosa. Mackenzie & Bush. Prairie Grove
and Farmington, Washington Co. Rare and local
(Ralph Shreve).
Sisyrinchium albidum Raf. 25061 Hazen, Prairie
Co.; Fayetteville, Washington Co.
Sisyrinchium furcatum Bicknell. 25050 Hazen,
Prairie Co.
Sisyrinchium graminoides Bicknell. Fayetteville,
Washington Co. Common throughout the state.
Orchidaceae:
Cypripedium pubescens Willd. A specimen sent
from Red Star, Newton Co., compared with material
at the Herbarium of the New York Botanical Gar-
den was found to be typical for leaf and size; 24993
McNab, Hempstead Co.
106 Trans. Acad. Sci. of St. Louis
Spiranthes ovalis Lindl. (Ibidium ovale (Lindl.)
House). Conway, Faulkner Co.
Spiranthes vernalis Engelm. & Gray. Fayette-
ville, Washington Co.; 8146 Ozark, Franklin Co.
Corrallorrhiza Wisteriana Conrad. Fayetteville,
Washington Co.; Fulton, Hempstead Co.
Pogonia verticillata (Willd.) Nutt. 26894 Law-
rence, Garland Co.
DICOTYLEDONAE.
Salicaceae:
Populus balsamifera var. virginiana (Henry) Sarg.
Common Cottonwood. Formerly Populus deltoides
Marsh. (P. monilifera Ait.) Distribution general
along streams.
Salix cordata Muhl. J ohnston, Washington Co.
Salix nigra var. altissima Sarg. Tall black wil-
low. 5394 Fulton, Hempstead Co.
Saliaz longifolia Muehl. Sand Bar Willow. 24978,
McNab, Hempstead Co.; 25013, Little Rock, Pulaski
Co.; 25085, Helena, Phillips Co.
Salix longipes var. Wardii (Bebb) Schneider.
Ward’s willow. Fayetteville, Washington Co.;
Eureka Springs and Beaver, Carroll Co.; Cotter,
Marion Co.
Salix longipes var. Wardii x nigra. 20500 Eureka
Springs, Carroll Co.
Leitneriaceae:
Leitneria floridana Chapm, Cork tree. 4792
Moark, 6074 Corning, Clay Co.; St. Francis River
swamps (S. M. Coulter).
Supplement to Catalogue of Arkansas Plants 107
Juglandaceae:
Carya alba var. ficoides Sarg. 20990 and 22198
Van Buren, Crawford Co.
Carya alba var. ovoidea Sarg. 29441 McNab,
Hempstead Co.
Carya alba var. subcoriacea Sarg. The common
form in Southern and Eastern Arkansas.
Carya Buckleyi var. arkansana Sarg. 20991 Van
Buren, Crawford Co. (type locality) ; 20638 Fulton,
Hempstead Co.; 10524 Doddridge, Miller Co. Com-
mon throughout the state.
forma pachylemma Sarg. Fulton, Hempstead
Co. (type locality).
Carya cordiformis var. latifolia Sarg. 8219 Fay-
etteville, Washington Co.; 20671 Cotter, Marion Co.
Carya leiodermis Sarg. 8953 Fulton, Hempstead
Co.; 26653 Helena, Phillips Co.; 26491 Mt. Nebo,
Yell Co.; 26424 Magazine Mountain, Logan Co.
Carya ovalis var. obovalis Sarg. Northern Ar-
kansas.
Carya ovalis var. obcordata Sarg. 22267 Fulton,
Hempstead Co.
Carya texana DC. Bitter Pecan. Van Buren,
Crawford Co., bottom lands of Arkansas River (G.
M. Brown),
X Carya Brownii Sarg. Van Buren, Crawford
Co., bottom lands of Arkansas River. A hybrid be-
tween Carya cordiformis and C. pecan.
X Carya Brownii var. varians Sarg. Van Buren,
Crawford Co. (G. M. Brown).
Carya alba x Buckleyi arkansana. 20989, 22210
Van Buren, Crawford Co.
108 Trans. Acad. Sci. of St. Lowis
Betulaceae:
Corylus rostrata Ait. Beaked Hazelnut. 26692
Jonesboro, Craighead Co.
Fagaceae:
Castanea alnifolia var. floridana Sarg. 413 Me-
Nab, Hempstead Co.; 8440 Benton, Saline Co.;
26536 Hardinville, Faulkner Co. (C. Margaretta
Ashe).
Castanea ozarkensis Ashe. Ozark Chinquapin.
(C. arkansana Ashe). War Eagle, Madison Co.
(type locality); Benton, Washington, Carroll and
other northern and western counties. (Formerly re-
ferred to C. pumila.)
Fagus grandifolia var. caroliniana Fern. & Rehd.
Beech. 8076 Gum Springs, Clark Co.; Pettigrew,
Madison Co., and along Mulberry Creek in Madison
and Franklin Cos. The forma mollis Fern. & Rehd.
has also been found at Helena, Phillips Co.
Quercus arkansana Sarg. Arkansas Oak. 5383,
7188, 8043, ete. McNab, Hempstead Co. Locally com-
mon in sand hills northwest of Fulton.
Quercus borealis Michx. Coll. by W. W. Ashe in
Ozark National Forest near Lurton, Newton Co.
Quercus borealis var. maxima Ashe. Northern
Red Oak. (Formerly erroneously called Q. rubra.)
6360 Beaver, 20510 Eureka Springs, Carroll Co.;
14300 Cotter, Marion Co.; Conway, Faulkner Co.;
Fayetteville and Savoy, Washington Co., and in all
of the northern, western and central counties.
Supplement to Catalogue of Arkansas Plants 109
Quercus coccinea Wang. Scarlet Oak. Jones-
boro, Craighead Co.; reported by R. M. Harper
from near Forrest City, St. Francis Co.
Quercus Durandii Buckley. Durand’s Oak. 5378,
6818, etc. Fulton, 8050 McNab, Hempstead Co.
Quercus nigra var. heterophylla (Ait.) ‘Ashe.
23995 Texarkana, Miller Co.; 24011 McNab, Hemp-
stead Co.; 24312 Shirley, Van Buren Co.
Quercus palustris Muench. Pin Oak. 6069 Corning,
Clay Co.; 21000 Van Buren, Crawford Co.; Conway,
Faulkner Co.; Russellville, Pope Co.; Clarendon,
Monroe Co. Reported by Harvey but omitted by
Branner & Coville.
Quercus Phellos L. Willow Oak. Conway, Faulk-
ner Co.; Pine Bluff, Jefferson Co.; ‘Alma, Crawford
Co.; Clarksville, Johnson Co. Reported by Harvey
but omitted by Branner & Coville.
Quercus prinoides Willd. Shin Oak, Prairie Oak.
Benton Co., margins of prairies (F. L. Harvey).
Quercus obtusa Ashe. Water Oak. Southern
Arkansas. Q. rhombica Sarg.)
Quercus rubra L. Southern Red Oak. (Formerly
Quercus triloba Michx., Q. digitata Sudw., Q. falcata
Michx.) Common throughout except in parts of
Ozark region.
Quercus rubra var. leucophylla Ashe. Little
Rock, Pulaski Co.; Fulton, Hempstead Co.; Stutt-
gart, Arkansas Co.; Conway, Faulkner Co.
Quercus rubra var. pagodaefolia (Ell.) Ashe.
Coastal Plains, S. E. Arkansas (fide Roland M.
Harper).
110
Trans. Acad. Sci. of St. Louis
Quercus Shumardu Buckley, Spotted Oak. (For-
merly reported as Quercus texana.) Fulton, 20635
McNab, Hempstead Co.; Dermott, Chicot Co.; Sa-
voy, Washington Co.; 6009 Cotter, Marion Co.
Quercus Shumardii var. Schneckii (Britton) Sarg.
Spotted Oak. 8182 Fayetteville, Washington Co.;
Benton Co.; Madison Co.; 4453 Eureka Springs,
Carroll Co.; 8949 Fulton, Hempstead Co.
Quercus stellata Wang. Post Oak. (Quercus minor
Sarg.) General distribution. Listed by Harvey but
omitted by Branner and Coville.
Quercus stellata var. araniosa Sarg. Sand Post
Oak. 22454, 8985, Texarkana, Miller Co.; 8439 Ben-
ton, Saline Co.
Quercus stellata var. Margaretta Ashe. Southern
Arkansas.
Quercus velutina var. missouriensis Sarg. Ben-
ton Co.; Washington Co.; Austin, Lonoke Co.; Con-
way, Faulkner Co.
Hybrid Oaks:
X Quercus Bushii Sarg. (Q. marilandica x Q.
velutina.) 20509 Eureka Springs, Carroll Co.;
Fayetteville, Washington Co. (Mock St. and also on
Lindell Ave.)
X Quercus Rudkinii Britton. (Q. marilandica x Q.
Phellos.) 10504, 23999, ete. Fulton, Hempstead Co.
X Quercus Schochiana Dieck. (Q. palustris x Q.
Phellos.) 27096 Dover, Conway Co.; Faulkner Co.
(D. Demaree),
Supplement to Catalogue of Arkansas Plants 111
X Quercus subfalcata Trel. (Q. Phellos x Q.
rubra.) 22466, 24013, etc. Fulton and McNab, Hemp-
stead Co.; Cove Creek, Faulkner Co. (D. Demaree).
Quercus Durandii x stellata. 12659, 20715, ete. Mc-
Nab, Hempstead Co.
Quercus nigra x Shumardii. 22301 McNab, Hemp-
stead Co.
Quercus nigra x rubra. 23139, 24876, ete. Hot
Springs, Garland Co.
Ulmaceae:
Ulmus serotina Sarg. 6944, Jasper, Newton Co.;
21013 Van Buren, Crawford Co.; 26603 Magnet
Cove, Hot Springs Co.; 22277 McNab, Hempstead
Co.
Celtis laevigata var. texana Sarg. Southern
Hackberry. 412 McNab, Hempstead Co.; 20491
Eureka Springs, Carroll Co. Common throughout
the state.
Celtis occidentalis var. canina (Raf.) Sarg. Fay-
etteville, Goshen, Washington Co.; 14329 Cotter,
Marion Co.
Celtis occidentalis var. crassifolia. (Lam.) Gray.
Rough leaved Hackberry. 20672 Cotter, Marion
Co.; 22208, 21012 Van Buren, Crawford Co.
Celtis pumila var. georgiana (Small) Sarg. Savoy,
West Fork, Washington Co.
Moraceae:
Broussonetia papyrifera (L.) Vent. (P apyrius
papyrifera (lL). Kuntze). Cultivated, occasionally
escaped. Paper Mulberry.
412 =: Trans. Acad. Sct. of St. Louis
Aristolochiaceae :
Asarum reflezum. Bicknell. Johnson, Goshen,
Kessler Mountain, West Fork, Washington Oo.
Frequent in Ozark region.
Aristolochia reticulata. Nutt. 8058 McNab, Hemp- —
stead Co.; 8384 Ashdown, Little River Co. Reported
in Nuttall’s Arkansas Flora but omitted from all
subsequent lists.
Polygonaceae:
Eriogonum longifolium Nutt, 14319 Cotter, Marion
Co. In sandy soil.
Eriogonum hirsutum Nutt. North Arkansas and
Missouri border region, confined to caleareous
rocks.
Polygonum densiflorwm Meisn. (Persicaria per
toricensis Small.) St. Francis River, Mississipp!
Co. (S. M. Coulter).
Chenopodiaceae :
Chenopodium Botrys L. 14304 Cotter, Marion
Co.
Amaranthaceae:
Froelichia gracilis Moq. Fayetteville, 8215 Brent-
wood, Washington Co.
Nyctaginaceae:
Boerhaavia erecita L. 8065 Arkadelphia, Clark Co.;
24257 Hot Springs, Garland Co.
Supplement to Catalogue of Arkansas Plants 113
Caryophyllaceae:
Cerastium brachypodum (Engelm.) Robinson. Short
Stalked Chickweed. Fayetteville, Washington Co.
Waste fields.
Cerastium semidecandrum L. Fayetteville, Wash-
ington Co.
Stellaria media (L.) Cyrill. Common Chickweed.
Waste places over entire state, sometimes flowering
during all months of winter.
Saponaria Vaccaria L. Cow herb. Conway, Faulk-
ner Co., near tunnel (H. E. Wheeler).
Silene nivea (Nutt.) Otth. Fayetteville, Washington
Co., escaped from cultivation.
Portulacaceae:
Talinum calycinum Engelm, 4563 Eureka Springs,
Carroll Co.; 26215 Cotter, Baxter Co.
Talinwm parviflorum Nutt. Thin soil above
sandstone cap rock of waterfall near Winslow,
Washington Co.; Fourche Mountain, Little Rock,
on *‘Pulaskite’’ rocks.
Nymphaeaceae:
Cabomba caroliniana Gray. Near Blytheville,
Mississippi Co.; St. Francis River swamps (S. M.
Coulter).
Ranunculaceae:
Anemone caroliniana Walt. Fayetteville, Wash-
ington Co.; Conway, Faulkner Co.
114
Trans. Acad. Sci. of St. Louis
Anemone decapetala Ard. Along sandstone cliffs,
south slopes, near base of Magazine Mountain, Logan
Co.
Delphinium Nortonianum Mackenzie & Bush. Fay-
etteville, Washington Co.
Delphinium Treleasei Bush. 27038 Eureka Springs,
Carroll Co.
Delphinium Penardii Huth. 5585 Beaver, Car-
roll Co. Frequent in barrens and on prairies of
Ozark region.
Ranunculus hispidus Michx. 5597 Eureka Springs,
Carroll Co.
Thalictrum texanum (Gray) Small, 7151 Fulton,
Hempstead Co.
Clematis versicolor Small. (Viorna versicolor
Small.) 6352 Beaver, Carroll Co.; 8226 Westfork,
Fayetteville, Washington Co.
Clematis reticulata Walt. (Viorna reticulata Small.)
10508 Columbus, Hempstead Co.
Clematis paniculata Thunb. 20498 Eureka Springs,
Carroll Co. Introduced along rocky creek.
Anonaceae:
Asimina parviflora (Michx.) Dunal. Crowley’s
- Ridge north of Helena, Phillips Co. (fide Roland M.
Harper.)
Menispermaceae:
Cocculus carolinus (L.) DC. (Epibaterium car-
olinum Britton). 6005 Cotter, Marion Co.; 22193
Van Buren, Crawford Co.; Fayetteville, Washing-
ton Co. Common throughout the state.
Supplement to Catalogue of Arkansas Plants 115
Calycocarpum Lyoni (Pursh) Nutt. 25153 Helena,
Phillips Co.; 26541 Cove Creek, Faulkner Co.;
24290 Shirley, Van Buren Co.
Fumariaceae:
Corydalis campestris (Britton) new comb. (Cap-
noides campestre Britton). Benton Co. (Type
locality).
Cruciferae:
Coronopus didymus (L.) J. E. Sm. 24989 Fulton,
Hempstead Co.; Conway, Faulkner Oo. A trouble-
some pasture weed.
Sisymbrium altissimum L. (Norta altissima Brit-
ton). 8158 Ozark, Franklin Co.; Washington Co.
Erysimum inconspicuum (Wats.) MacM. North-
west Arkansas (HE. N. Plank).
Camelina sativa (L.) Crantz. Gold of Pleasure.
Fayetteville, Washington Co. Naturalized from
Europe.
Dentaria laciniata Muhl. 24855 Magazine Moun-
tain, Logan Co.
Cardamine pennsylvanica Muhl. 24458 Hot Springs,
Garland Co.; Fayetteville, Washington Co.
Cardamine parviflora L. 7176 Fulton, Hempstead
Co.; 24451 Hot Springs, Garland Co.; Conway,
Faulkner Co.; Fayetteville, Washington Co.
Droseraceae:
Drosera annua E. L. Reed. Conway, Faulkner Co.,
Campus of State Teachers College.
116
Trans. Acad. Sci. of St. Louis
Saxifragaceae:
Sazifraga texana Buckl. (Micranthes texana
Small). Conway, Faulkner Co.; Fayetteville,
Washington Co.
Heuchera arkansana Rydb. (May be the H. villosa
of Harvey.) On cliffs, Northwest Arkansas. .
Heuchera hirsuticaulis (Wheelock) Rydb. Shaded
north slopes, on Boone chert hills, Washington Co.
Heuchera macrorhiza Small. Fayetteville, Wash-
ington Co.
Heuchera parviflora Bartl, Springdale, Wash-
ington Co.; Shirley, Van Buren Co. On moist cliffs.
Flowers in October.
. Philadelphus pubescens Schrad. Cove Creek, Faulk-
ner Co.; 6934 Jasper, Newton Co.; 6033 Rush,
Marion Co.; Shirley, Van Buren Co.; Magazine
Mountain Logan Co.
Hydrangea cinerea Small. 6959 Heber Springs,
Cleburne Co.; 25160 Helena, Phillips Co.; 25205
Shirley, Van Buren Co.
Ribes Cynosbati L. Prickly Gooseberry. (Gross-
laria Cynosbati Mill.). 242 Rich Mountain, Polk Co.;
Magazine Mountain, Logan Co.
Kibes: missouriense Nutt. Gooseberry (Grossul-
aria missouriensis Coville & Brit.) 5572 Beaver,
Carroll Co.
Hamamelidaceae:
Hamamelis macrophylla Pursh. Witch-hazel. 249
Rich Mountain, Polk Co.; 8441 Benton, Saline Co.;
22296 McNab, Hempstead Co.
Supplement to Catalogue of Arkansas Plants 117
Hamamelis vernalis Sarg. Spring-blooming Witch-
hazel. 6032 Rush Marion Co.; 12649, Mena, Polk
Co.; Goshen, Washington Co. Common along rocky
streams of Ozark region.
Hamamelis vernalis var. tomentella Sarg. 20494
Eureka Springs, Carroll Co.
Rosaceae:
Physocarpus intermedius (Rydb.) Schneider. Nine
Bark. (Opulaster intermedius Rydb.) 4385 Eureka
Springs, Carroll Co.; 5573 Beaver, Carroll Co..; 5982
Cotter, Marion Co. Probably the Neillia opulifolia
Benth. & Hook., observed by Lesquereux and by Har-
vey, belongs here.
Malus angustifolia (Ait.) Michx. Narrow-leaved
Crabapple. 20719 Arkadelphia, Clark Co.
Malus iowensis var. Palmeri Rehd. 10581 Wash-
ington, Hempstead Co.
Crataegus. Crus-galli group:
Crataegus bellica Sarg. 5335, 5336 Fulton, Hemp-
stead Co, (type locality).
Crataegus Bushii Sarg. 5344 Fulton, Hempstead
Co. (type locality); 277 Rich Mountain, Polk
County; 24531 High Point, Garland Co. €
Crataegus erecta Sarg. 21017 Van Buren, Craw-
ford County.
Crataegus Palmeri Sarg. Fayetteville, Washington
Co.
Crataegus paradoxa Sarg. 20993 Van Buren, Craw-
ford Co.
118
Trans. Acad. Sci. of St. Louis
Crataegus pilifera Sarg. 5540 Eureka Springs,
Carroll Co.; Van Buren, Crawford Co.
Crataegus subpilosa Sarg. Eureka Springs, Car-
roll Co. (type locality).
Crataegus triwmphalis Sarg. 5371, 6825, Fulton,
Hempstead Co. (type locality).
Crataegus palliata Sarg. Fulton, Hempstead Co.
(type locality).
Crataegus villiflora Sarg. 5519, 5522. Eureka
Springs, Carroll Co.; 20992, 22205 Van Buren,
Crawford Co. |
Crataegus. Punctatae group:
Crataegus fastosa Sarg. Fulton, Hempstead Co.
(type locality).
Crataegus secta Sarg. 19028 Sulphur Springs,
Benton Co.
Crataegus sordida var. villosa Sarg. 20709, 22266
McNab, Hempstead Co.
Crataegus sucida Sarg. 4387 Eureka Springs,
Carroll Co.; 8179 Fayetteville, Washington Co.
Crataegus verruculosa Sarg. 24453 Hot Springs,
Garland Co.
Crataegus. Virides group:
Crataegus amicalis Sarg. Fulton, Hempstead Co.,
(type locality).
Crataegus blanda Sarg. 20712 McNab, Hempstead
Co.; 24496 Hot Springs, Garland Co.
Crataegus micrantha Sarg. Fulton, Hempstead
Co. (type locality).
Supplement to Catalogue of Arkansas Plants 119
Crataegus velutina Sarg. Fulton, Hempstead
Co. (type locality).
Crataegus viridis L. 4789 Moark, Clay Co.; 5345
Fulton, Hempstead Co.; 8134 Little Rock, Pulaski
Co.; 8221 Greenland, Washington Co. The collec-
tions credited to Harvey as C. arborescens Ell. prob-
ably belong here, since this is the species occurring
in the Fayetteville region.
Crataegus. Intricatae group:
Crataegus Harveyana Sarg. Eureka Springs, Car-
roll Co. (type locality); 26849 Hot Springs, Gar-
land Co.
Crataegus leioclada Sarg. 4388, 5539, 5547, Eureka
Springs, Carroll Co.
Crataegus neobushii Sarg. 5546 Eureka Springs,
Carroll Co.
Crataegus villicarpa a 5525 Eureka Springs,
Carroll Co.
Crataegus padifolia var. incarnata Sarg. 6026
Cotter, Marion Co. ; 26611 Magnet Cove, Hot Springs
Co.
Crataegus. Tenuifoliae (?) group:
Crataegus lacera Sarg. Fulton, Hempstead Co.
(type locality).
Crataegus. Molles group:
Crataegus arkansana Sarg. Type (at Arnold
Arboretum) from seed collected at Newport, Ar-
kansas,
120 Trans. Acad. Sci. of St. Louis
Crataegus brachyphylla Sarg. 432, 8975, 9393,
9392, 10607, 16333, 20711, McNab, Hempstead Co.
(type locality).
Crataegus invisa Sarg. 5397, Fulton (type lo-
cality), 6879, 7193, 7196, McNab, Hempstead Co.
Crataegus induta Sarg. Fulton, Hempstead Co.
(type locality).
Crataegus limaria Sarg. a Hempstead Co.
(type locality).
Cartaegus transmississippiensis Sarg. 8419 Cot-
ter, Marion Co. (type locality).
Crataegus. Dilatatae group:
Crataegus coccinoides Ashe. Farmington, Washing-
ton Co.
Crataegus. Uniflorae group:
Crataegus trianthophora Sarg. 6970 Heber Springs,
Cleburne Co.; 26535 Cove Creek, Faulkner Co.;
26885 Lawrence, Garland Co.; 14322 Cotter,
Marion Co.; Fulton, Hempstead Co. (B. F. Bush
5690) ; Baker Springs, Howard Co. (J. H. Kellogg).
Crataegus. Brachyacanthae group:
Crataegus brachyacantha Sarg. & Engelm. 315
Texarkana, Miller Co.; 8386 Ashdown, Little River
Co.
Crataegus. Macracanthae group:
Crataegus carrollensis Sarg. Eureka Springs,
Carroll Co. (type locality).
Crataegus hispidula Sarg. 5542 Eureka Springs,
Carroll Co.
Supplement to Catalogue of Arkansas Plants 121
Crataegus mollicula Sarg. 6023 Rush, Marion Co.
Crataegus pudens Sarg. 20998, 22204 Van Buren,
Crawford Co.
Crataegus tomentosa lL. This species, reported
by Branner & Coville, is found in ‘Arkansas, but
the variety punctata Gray (C. punctata Jacquin),
reported also by them, probably does not occur
here.
Crataegus aestivalis (Walt.) T. & G., C. coccinea
L., C. flexispina (Moench) Sarg. and C. subvillosa
Schrad., listed for Arkansas by Branner & Coville,
very probably are not found here and should be
dropped from the state plant list.
C. cordata Ait. (=C. phaenopyrum (L. f.) Medic.)
and C. crus-galli L. reported by Harvey may occur
in the northern counties.
Crataegus hybrids: :
X Crataegus notha Sarg. A hybrid between C.
apifolia and C. brachyphylla. 8974, 20646, 20716
McNab, Hempstead Co.
Alchemilla arvensis Scop. Lady’s Mantle. 23133
Hot Springs, Garland Co.
Potentilla recta L. Fayetteville, Washington Co.
Prunus mexicana Watson. Mt. Nebo, Yell Co.;
Rich Mountain, Polk Co.; 7149 Fulton, 16329 McNab,
Hempstead Co.; 20474 Eureka Springs, Carroll Co.;
20668 Cotter, Marion Co.
Prunus mecicana var. fultonensis Sarg. 12663 Ful-
ton, Hempstead Co. (type locality).
122
Trans. Acad. Sci. of St. Louis
Prunus mexicana var. polyandra Sarg. Fulton,
Hempstead Co.
Prunus Munsoniana Wight & Hedrick. Fulton,
Hempstead Co.
Prunus wnbellata Ell. 8109 Gifford, Hot Springs Co. ;
10597 Washington, Hempstead Co.
Prunus umbellata var. tarda (Sarg.) Wight. 105395
Arkadelphia, Clark Co.
Prunus angustifolia var. varians Wight. 7148 Ful-
ton, Hempstead Co.
Rosa Eglanteria L. (Rosa rubiginosa L.) Escaped
from cultivation, Fayetteville, Washington Co.
Rosa Lyonii Pursh. 24957 Hot Springs, Garland
Co.; 24994 McNab, Hempstead Co.; 25055 Mesa,
Prairie Co.; 25103 Helena, Phillips Co.
Rosa Palmeri Rydb. 8067 Ashdown, Little River
Co.
Rosa setigera var. tomentosa T. & G. 22294 Mc-
Nab, Hempstead Co. ; 5978 Cotter, Marion Co. ; 20507
Eureka Springs, Carroll Co.
Rosa subserrulata Rydb. 8159 Ozark, Franklin
Co. ; 10531 Ft. Lynn, Miller Co.; 6960 Heber Springs,
Cleburne Co.; 8137 Little Rock, Pulaski Co.
Rosa texarkana Rydb. Type collected by Eggert,
Texarkana, Miller Co.
Rubus occidentalis L. 5574 Beaver, Carroll Co.
Common throughout Ozark region.
Rubus ostryaefolius Rydb. 16315 Texarkana, Mil-
ler Co.
Supplement to Catalogue of Arkansas Plants 123
Rubus Andrewsianus Blanchard. Fulton, Hemp-
stead Co. The common high blackberry throughout
the State.
Rubus rubrisetus Rydb. 327 Texarkana, Miller Co.;
20727 Gum Springs, Clark Co.
Leguminosae:
Amorpha croceolanata Watson. (A. fruticosa var.
croceolanata Schneider). 948 Cove Creek, Faulkner
Co.; 23051, 26863 Hot Springs, Garland Co.; 26918
Magnet Cove, Hot Springs Co.
Amorpha nitens Boynton. 23148, 26862 Hot Springs,
Garland Co.; 26915 Magnet Cove, Hot Springs
County.
Amorpha tennesseensis Shuttl. 20659 Cotter,
Marion Co.; 24251 Hot Springs, Garland Co.; Van
Buren, Crawford Co. (G. M. Brown).
Amorpha glabra Desf. 24187 Magazine Moun-
tain, Logan Co.; Hot Springs, Garland Co.; Rich
Mountain, Polk County.
Baptisia sulphurea Emgelm. Little Rock, Pu-
laski Co. (H. E. Wheeler).
Cassia Medsgeri Shafer. Fayetteville and West-
fork, Washington Co.; 6885 Eureka Springs, Car-
roll Co.; Cotter, Marion Co. Frequent in Ozark
region and perhaps throughout the state.
Cladrastis lutea (Michx.) K. Koch. Kessler
Mountain, near Farmington, Washington Co.; Mag-
azine Mountain, Logan Co.; Shirley, Van Buren
Co.; 8408 Cotter, Marion Co.
124
Trans. Acad, Sci. of St. Louis
Lathyrus latifolius L. Perennial Pea. Fayette-
ville, Washington Co. Escaped from cultivation.
Lespedeza procumbens Michx. Fayetteville, Wash-
ington Co.; Eureka Springs, Carroll Co.
Medicago arabica Huds. Spotted Medick. Fay-
etteville, Washington Co. (R. H. Austin). Escaped
from cultivation.
Medicago lupulina L. Black Medick. Fayette-
ville, Washington Co. Introduced.
Melilotus officinalis (L.) Lam. Yellow Sweet
Clover. Washington Co., waste places, introduced
and becoming common in many parts of the state.
Melilotus alba Desv. White Sweet Clover. In-
troduced and becoming abundant all over the state.
Desmodium paniculatum var. pubens T. & G.
4475 Eureka Springs, Carroll Co.
Petalostemum villosum Nutt. 5994 Cotter, Marion
Co.
Phaseolus polystachyus (l.) B. S. P. Wild Kid-
ney Bean. 8396 Cotter, Marion Co.; 10592 Wash-
ington, Hempstead Co.
Sesbania macrocarpa Nutt. Fulton, Hempstead
Co. (4405 J. M. Greenman).
Stizolobium Deeringianuwm Bort. Velvet Bean.
McNab, Hempstead Co. (4436 J. M. Greenman).
Wistaria macrostachya Nutt. Fulton and Me-
Nab, Hempstead Co.
Strophostyles umbellata (Muhl.) Britton. 12659
McNab, Hempstead Co.
Supplement to Catalogue of Arkansas Plants 125
Strophostyles helvola (L.) Britton. Fayette-
ville, Washington Co.
Trifolium hybridum L. Alsike Clover. Fayette-
ville, Washington Co. Introduced. (R. H. Austin).
Trifolium dubium Sibth. Fayetteville, Washing-
ton Co.; Eagle Mills, Ouachita Co. Introduced.
Vicia angustifolia (L.) Richard. 24510 Hot Springs,
Garland Co. Introduced.
Vicia sativa L. Fayetteville, Washington Co.
Introduced.
Vicia tetrasperma (L.) Moench. 5395 Fulton,
Hempstead Co. Introduced.
Geraniaceae:
Erodium cicutarium (L.) L’Hér. University of
Arkansas campus, Fayetteville (A. D. Oxley).
Geranium pusillum Burm, f. University of Ar-
kansas campus, Fayetteville.
Oxalidaceae:
Oxalis Brittonae Small. England, Lonoke Co.
Oxalis texana (Small) Fedde. Introduced in
favored places or as weed in greenhouses,
Oxalis Langliosii (Small) Fedde. Fayetteville,
Washington Co.
Oxalis interior (Small) Fedde. Benton Co. (type
locality) (E. N. Plank).
Linaceae:
Linum medium (Planch.) Small. Hazen, Prairie
Co. (Roland M. Harper).
126 Trans. Acad. Sci. of St. Louis
Zygophyllaceae:
Tribulus terrestris L. Ft. Smith, Sebastian Co.;
6094 Corning, Clay Co.
Rutaceae:
Ptelea polyadenia Greene. 5923, 5921 Cotter, Marion
Co.; 6957 Heber Springs, Cleburne Co.
Zanthoxyllum americanum Mill. Savoy, Wash-
ington Co.
Meliaceae:
Melia Azedarach L. Escaped from cultivation,
Fulton, Hempstead Co.
Euphorbiaceae:
Andrachne phyllanthoides (Nutt.) Muell. Arg.
5528 Eureka Springs, 27053 Berryville, Carroll Co.;
Magazine Mtn., Logan Co.; Hot Springs, Garland
Co.
Croton Engelmanni var. albinoides Ferg. Ful
ton, Hempstead Co.; Texarkana, Miller Co.; Ash-
down, Little River Co.
Crotonopsis elliptica Willd. 6962 Heber Springs,
Cleburne Co., also common in barrens of Ozark re-
gion.
Euphorbia arkansana Engelm. & Gray. (Tithy-
malus arkansanus Kl. & Garcke). Clark Co. (C.
Woolsey).
Euphorbia commutata Engelm. (Tithymalus com-
mutatus Kl. & Garecke). Eureka Springs, Car-
roll Co. (C. Woolsey).
Supplement to Catalogue of Arkansas Plants 127
Euphorbia Nuttallii Engelm. (Chamaesyce Nuttallii
Small). 6333 Beaver, Carroll Co.
Tragia ramosa Torr. 4382 Eureka Springs,
Carroll Co.
Anacardiaceae:
Cotinus americanus Nutt. American Smoke-tree,
Chittamwood. 5971, 10556 Cotter, Marion Co.;
Van Buren, Crawford Co. (G. M. Brown).
Rhus trilobata Nutt. Polecat-bush. (Schmaltzia
trilobata Small). 4384 Eureka Springs, Carroll
Co.; 4754, 8411 Cotter, Marion Co. Common on
rocky bluffs of Ozark region. Schmaltzia serotina
Greene also occurs in the northern part of the state
and may be distinct.
Rhus quercifolia (Michx.) Steud. Poison Oak.
258 Rich Mountain, Polk Co.; 24810 Blue Mountain,
Logan Co.; 25052 Hazen, Prairie Co.
Aquifoliaceae:
Ilex caroliniana (Walt.) Trel. 12661, 14648 Mc-
Nab, Hempstead Co.; 23060 Hot Springs, Garland
Co.; Cove Creek, Faulkner Co.
Aceraceae:
Acer floridanum (Chapm.) Pax. McNab, Hemp-
stead Co.; Crowley’s Ridge (Roland M. Harper).
Acer leucoderme Small. 24508, 26474, ete. Hot
Springs, Garland Co.; Baker Springs, Howard Co.
(J. H. Kellogg).
Acer saccharum var. glaucum (Pax.) Sarg. 14313
Cotter, Marion Co.; 20488 Eureka Springs, Carroll
Co. The common Sugar Maple of the state.
128
Trans. Acad. Sci. of St. Louis
Acer saccharum var. Rugellii (Pax.) Rehd. Eureka
Springs, Carroll Co.; near Hindsville, Madison Co.
Acer rubrum var. Drummondii (Hook. & Arn.)
Sarg. (Acer Drummondii Hook. & Arn.) 7164 Ful-
ton, Hempstead Co.; St. Francis River swamps (S.
M. Coulter).
Acer rubrum var. tridens Wood. 8364 Ashdown
Little River Co.; Fulton, Hempstead Co.
Acer Negundo var. texanum Pax. 6977 Heber
Springs, Cleburne Co.; 8072 Arkadelphia, Clark Co.;
14639 Cotter, Marion Co.; 22261 Fulton, Hempstead
Co. The form latifolia Sarg. is also found at Fulton.
Hypocastanaceae:
Aesculus discolor var. mollis Sarg. Red-flowered
Buckeye. (Aesculus austrina Small). 8078 Gum
Springs, Clark Co.: Austin, Lonoke Co.; Fulton,
Hempstead Co. Common over the state except in
parts of the Ozark region. :
Aesculus glabra var. micrantha Sarg. Fulton,
Hempstead Co. (type locality).
Aesculus glabra var. pallida Kirch. 8203 Wins-
low, Washington Co.
Aesculus glabra var. leucodermis Sarg. (Reported
by Harvey as A. glabra). Fayetteville, Washington
Co.; 4442 Eureka Springs, Carroll Co.
X Aesculus Bushii Schneider. Fulton, Hemp-
stead Co. (type locality). (B. F. Bush). A supposed
hybrid between A. glabra var. leiodermis and
A, discolor var. mollis.
Supplement to Catalogue of Arkansas Plants 129
Sapindaceae:
Sapindus Drummondit Hook & Arn. 5949 Cot-
ter, Marion Co.; Fulton, Hempstead Oo.; Savoy,
Washington Co.
Rhamnaceae:
Ceanothus ovatus Desf. 21015 Van Buren, Crawford
Co.; Farmington, Washington Co.
Ceanothus pubescens (T. & G.) Rydb. North Arkan-
sas, exposed limestone ledges.
Rhamnus lanceolata Pursh. 5931, (17239) Cotter,
Marion Co.; 20495 Eureka Springs, Carroll Co.
Vitaceae:
Ampelopsis cordata Michx. False Grape. Fayette-
ville, Washington Co.; 5933 Cotter, Marion Co.
Parthenocissus quinquefolia var. hirsuta (Don.)
Planch. 216 Huntsville, Madison Co.; 22189, 22209.
Van Buren, Crawford Co.
Parthenocissus quinquefolia var. Saint-Paulii
(Koehne & Graebn.) Rehd. 5909 Cotter, Marion Co. ;
22269 Fulton, Hempstead Co.
Vitis cinera Engelm. Winter Grape. 5932 Cotter,
Marion Co. Common throughout state.
Vitis Linsecomii var. glauca Munson. Post Oak
Grape. 5554 Eureka Springs, Carroll Co.; 22191
Van Buren, Crawford Co.; 228 MeNab, Hempstead
Co.; Fayetteville, Washington Co.
Vitis rotundifolia Michx. Muscadine. Conway,
Faulkner Co.; 257 Rich Mountain, Polk Co.; For-
rest City, St. Francis Co.; 6839 Columbus, Hemp-
stead Co.; 8470 Gifford, Hot Springs Co.; 6956
Heber Springs, Cleburne Co.
130 Trans. Acad. Sci. of St. Louis
Vitis palmata Vahl. 6052 Corning, Clay Co.;
10526 Ft. Lynn, Miller Co.; 12667 McNab, Hemp-
stead Co.
Tiliaceae:
Tila caroliniana Mill. 16336 McNab, Hempstead Co.
Tilia caroliniana var. rhoophila Sarg. 8074 Gum
Springs, Clark Co.
Tilia floridana Small. (Tilia pubesceus Ait., re-
ported by Coville, may belong here). 7204, 9401 Me-
Nab, Hempstead Co.; 20512 Eureka Springs, Car-
roll Co.; 20657 Cotter, Marion Co.; 20729 Gum-
Springs, Clark Co.
Tilia floridana var. hypoleuca Sarg. 5943 Cot-
ter, Marion Co.; Kessler Mountain near Farming-
ton, Washington Co.
Tilia nuda Sarg. Fulton, Hempstead Co.
Malvaceae:
Callirhoe alcaeoides (Michx.) Gray. University of
Arkansas campus, Fayetteville.
Callirhoe Bushii Fernald. Baldwin, Savoy, Wash-
ington Co.
Callirhoe triangulata (Leavenworth) Gray. 5977
Cotter, Marion Co.
Hibiscus incanus Wendl. 466, 16321 Fulton, Hemp-
stead Co.
Hibiscus lasiocarpus Cav. 8365 Ashdown, Little
River Co.; 26638 Helena, Phillips Co.
Malvastrum angustatum Gray. 6910 Harrison,
Boone Co.
Supplement to Catalogue of Arkansas Plants 131
Hypericaceae:
Ascyrum multicaule Michx. 22289 McNab, Hemp-
stead Co.
Hypericum apocynifolium Small. Hazen, Prairie
Co. and §. E. coastal plain. (Roland M. Harper).
Hypericum pseudomaculatum Bush. Fayetteville,
Washington Co.; 5957 Cotter, Baxter Co.; 8155
Ozark, Franklin Co.
Hypericum oklahomense E. J. Palmer. Cove Creek,
Faulkner Co.
Cistaceae:
Lechea villosa Ell. 8122 Malvern, Hot Springs
Co.
Helianthemum rosmarinifolium Pursh. 8162 Piney,
Johnson Co.
Violaceae:
Viola eriocarpa Schwein. Johnson and Elm Springs,
Washington Co. (V. pubescens Ait, of Branner and
Coville, probably belongs here).
Viola fimbriatula J. FE. Sm. 420 and 7153 Me-
Nab, Hempstead Co.
Viola Lovelliana Brainerd. Mena, Polk Co. (HK.
Brainerd). :
Viola missouriensis Greene. Osage Creek, Wash-
ington Co.; Benton Co.; 5393, 7165, Fulton, Hemp-
stead Co.
Viola primulifolia L. Washington, Hempstead
Co.; West Mountain near Hot Springs, Garland Co.
Viola sororia Willd. Fayetteville, Washington
Co.; 4782 Moark, Clay Co.
132
Trans. Acad. Sci. of St. Louis
Viola triloba Schwein. (This may be Harvey’s
‘var. cordata). Mt. Comfort, Washington Co.; 5374,
7150, 9390 Fulton, Hempstead Co. |
Viola triloba var. dilatata (Ell.) Brainerd. Mena,
Polk Co., West Fork, Washington Co.; (Westville,
Okla. E. Brainerd).
Viola papilionacea Pursh. 24465 Hot Springs,
Garland Co.; Fayetteville, Washington Co.; Mena,
Polk Co, (KE. Brainerd).
Viola striata Ait. Fayetteville, Washington Co.
Viola viarum Pollard. Along White River. (H.
Brainerd).
Lythraceae:
Cuphea petiolata (L.) Koehne, (Parsonsia petiolata
Rusby). 4423 Eureka Springs, Carroll Co.
Onagraceae:
Oenothera linearis Michx. (Kneiffia linearis Spach).
Fayetteville, Washington Co.; 23218 Top of Maga-
zine Mountain, Logan Co.; 5404 Fulton, Hemp-
stead Co.
Oenothera missouriensis Sims. (Megapterium mis-
souriense Spach). Eureka Springs, Carroll Co.
Oenothera arenicola (Small) Coker. (Kneiffia arent-
cola Small). 25038 Hazen, Prairie Co.
Cornaceae:
Cornus alternifolia L. Sylamore, Stone Co. (W.
W. Ashe), also in Washington Co.
Cornus stricta Lam. 6090 Corning, 10591 Moark,
Clay Co.; 12649a Mena, Polk Co.; 8363 Ashdown,
Little River Co.; 16324 Fulton, Hempstead Co.
«
Supplement to Catalogue of Arkansas Plants 133
Umbelliferae:
Ammoselinum Butleri (Engelm.) Coult. & Rose.
Credited to State in Coulter & Rose, Monograph of
Umbelliferae.
Angelica villosa (Walt.) B. S. P. 6024 Rush,
Marion Co.
Chaerophyllum texanum Coult. & Rose. 5409 Fulton,
Hempstead Co.
Daucus Carota L. Along roadsides and railroads,
Fayetteville, Washington Co.
Erigenia bulbosa (Michx.) Nutt. West Fork,
Washington Co.
Eryngium prostratum Nutt. 6048 Corning, Clay
Co.; 8166 London, Pope Co.
Hydrocotyle bonariensis Lam. Coastal Plain of
Arkansas. (Roland M. Harper).
Ptilimnium Nuttallii (DC.) Britton. 8027 Ful-
ton, Hempstead Co.; 8164 Piney, Johnson Co.
Ptilimnium capillaceum (Michx.) Raf. 26682
Jonesboro, Craighead Co.
Sium suave Walt. (Sium cicutaefolium Schrank).
St. Francis River swamps. (S. M. Coulter).
Torilis Anthriscus (L.) Bernh. Fayetteville, Wash-
ington Co. (Miss C. Chandler).
Zizia aurea (L.) Koch. 5559 Eureka Springs,
Carroll Co.
Zizia cordata (Walt.) DC. 5557 Eureka Springs,
Carroll Co.
134
Trans. Acad. Sci. of St. Lows
SYMPETALAE.,
Ericaceae:
Monotropa Hypopitys L. (Hypopitys americana
Small). Prarie View, Washington Co., densely
wooded north slopes of chert hills.
Rhododendron oblongifolium (Small) Millais. Aza-
lea, Honeysuckle. (Azalea oblongifolia Small).
6932 Jasper, Newton Co.; 6973 Heber Springs, Cle-
burne County; 10505 Columbus, Hempstead Co.;
10515 Doddridge, Miller Co.
Rhododendron rosewm (Loisel.) Rehd. Azalea,
Honeysuckle. 260 Rich Mountain, Polk Co.; 26466
Meaford, Garland Co.; Batesville, Independence Co. ;
Cove Creek, Faulkner Co.
Rhododendron canescens (Michx.) G. Don (Azalea
canescens Michx.). Yellville, Marion Co.
Vaccinium arboreum var. glaucescens (Greene)
Sarg. (Batodendron glaucescens Greene). 6942
Jasper, Newton Co.
Vaccinium vacillans Kalm. Fulton, Hempstead
Co. and occasional through Ozark region, but much
less common than the following variety.
Vaccinium vacillans var. crinitum Fernald. 5556 Eu-
reka Springs, Carroll Co.; 5989 Cotter, Marion Co.
Common throughout Ozark region.
Primulaceae:
Dodecatheon brachycarpa Small. Wheeler, Wash-
ington Co.
Steironema intermedium Kearney. Fayetteville,
Washington Co.
Supplement to Catalogue of Arkansas Plants 135
Sapotaceae:
Bumelia lycioides (L.) Pers. 25137, 26649 Helena,
Phillips Co.; 26725 McNab, Hempstead Co.
Ebenaceae:
Diospyros virginiana var. platycarpa Sarg. Large-
fruited Persimmon. 20669 Cotter, Marion OCo.,
also in northwestern counties.
Styracaceae:
Halesia monticola var. vestita Sarg. Silver-bell
Tree. 255 Rich Mountain, Polk Co.; 6978 Heber
Springs, Cleburne Co.; 24926 Hot Springs, Garland
Co.
Oleaceae:
Forestiera acuminata var. vestita E. J. Palmer.
Miller Co. (type by Bush); Fulton, Hempstead Co. ;
Van Buren, Crawford Co.
Ligustrum vulgare L. Privet. 14321 Cotter, Marion
Co. Escaped from cultivation.
Fraxinus pennsylvanica var. lanceolata (Borck.)
Sarg. Green Ash. 6073 Corning, Clay Co.; 5942
Cotter, Marion Co.; 8080 Arkadelphia, Clark Co.;
Fayetteville, Washington Co. Common along
streams throughout state.
Fraxinus profunda Bush. Pumpkin Ash. 6067
Corning, Clay Co.; St. Francis River swamps (S.
M. Coulter),
According to Harvey, Fravinus nigra Marsh (P.
sambucifolia L.) is found in ‘Arkansas. This is prob-
ably an error which has been repeated by others.
136 Trans. Acad, Sci. of St. Louis
No specimens are to be found in any of the Amer-
ican herbaria and Harvey left no specimens in the
University of Arkansas collections.
Gentianaceae:
Bartonia virginica (L.) B. S. P. 10586 Washing-
ton, Hempstead Co.; 26476 Lonsdale, Garland Co.
Centaurium texense (Griseb.) Fernald. (Erythraea
texensis Griseb. Northern Arkansas border counties.
Gentiana flavida Gray. (Dasystephena flavida
Britton.) Magazine Mountain, Logan Co.
Apocynaceae:
Amsonia ludoviciana Vail. Fayetteville, Wash-
ington Co.; Lonoke, Lonoke Co.
Apocynum cannabinum var. pubescens (R. Br.)
DC. 6092 Corning, Clay Co.
Vinca minor L, Periwinkle or Myrtle. Wash-
ington Co. Escaped locally from cultivation.
Asclepiadaceae : .
Asclepias stenophylla Gray. 8397 Cotter, Marion Co.
Asclepias Sullivantii Engelm. Fayetteville, Wash-
ington Co.
Asclepias perennis Walt. Corning, Clay Co.
Asclepias purpurascens L. 12648 Mena, Polk Co.
Acerates floridana (Lam.) Hitche. 25018 Hazen,
Prairie Co.
Convolvulaceae:
Cuscuta cuspidata Engelm. 16322 Fulton, Hemp-
stead Co.
Stylisma humistrata (Walt.) Chapman. 8111 Gif-
ford, Hot Springs Co. :
Supplement to Catalogue of Arkansas Plants 137
Polemoniaceae:
Phlox bifida Beck. War Eagle Creek near Hunts-
ville, Madison Co.
Hydrophyllaceae:
Nemophila phacelioides Nutt. Wheeler and Johnson,
Washington Co.; Hot Springs, Garland Co.
Boraginaceae:
Onosmodium occidentale Mackenzie. 8039 Fulton,
Hempstead Co.; Baxter Mountain, Washington Co.
Onosmodium subsetosum Mack. & Bush. 6041a
Northfork, Baxter Co.
Verbenaceae:
Verbena Drummondii (Lindl.) Baxt. Savoy, Wash-
ington Co.
Verbena Lambertit Sims. Son’s Chapel, Wash-
ington Co.
Labiatae:
Satureja glabella (Michx.) Briquet. (Clinopodium
glabellum Kuntze). West Fork, Washington Co., on
calcareous outcrops.
Lamium purpureum L. Fayetteville, Washington
Co. Introduced.
Monarda citriodora Cerv. Sulphur Springs, Ben-
ton Co.
Perilla frutescens (L.) Britton. 4437 Eureka
Springs, Carroll Co.; Monte Ne, Benton Co.; Crow-
ley’s Ridge, near Forrest City, St. Francis Co.
(Roland M. Harper).
138 Trans. Acad. Sci. of St. Lowis
Salvia azurea Lam. 451 McNab, Hempstead Co.
Salvia lanceifolia Poir. 4485 Eureka Springs, Car-
roll Co.
Solanaceae:
Datura Metel L. Egger, Polk Co. (R. H. Rosen).
Introduced.
Physalis missouriensis Mack. & Bush. 6955 Fulton,
Hempstead Co.
Bouchetia anomala (Miers) Britt. & Rusby. Sum-
mers, Washington Co., prairie glades.
Solanum elaeagnifolium Cav. 338 Texarkana, Miller
Co.; Fayetteville, Washington Co.
Solanum Torreyi Gray. 23009 Pinnacle, Pulaski Co.
Serophulariaceae:
Agalinis fasciculata (Ell.) Raf. 8464 Gifford,
Hot Springs Co.; 8472 Gum Springs, Clark Co.
Buchnera elongata Sw. Prairies of Arkansas,
coastal plain (Roland M. Harper).
Aureolaria grandiflora var. serrata (Torr.) Pen-
nell. (Dasystoma serrata Small) 10603 Washing-
ton, 8977 McNab, Hempstead Co.
LIinaria Cymbalaria (L.) Mill. Kenilworth Ivy.
Fayetteville, Washington Co. Occasionally escaped.
Paulownia tomentosa (Thunb.) Steud. 10527 Ft.
Lynn, Miller Co.; Monticello, Drew Co.; Hot
Springs, Garland Co. Escaped from cultivation.
Pentstemon arkansanus Pennell. 8143 Little Rock,
Pulaski Co.; 24556 Hot Springs, Garland Co.;
25167 Shirley, Van Buren Co.
Supplement to Catalogue of Arkansas Plants 139
Scrophularia neglecta Rydb. 12687 McNab, Hemp-
stead Co.
Veronica arvensis L. Fayetteville, Washington Co.
and throughout the state.
Veronica serpyllifolia L. Withrow Springs, Madi-
son Co.
Veronica Tournefortii C. C. Gmel. Fayetteville,
Washington Co., University of Arkansas campus.
Orobanchaceae:
Orobanche ludoviciana Nutt. (Myzorrhiza ludo-
viciana Rydb.) Kingdon Springs, Marion Co.
(Roy Morrow).
Rubiaceae:
Cephalanthus occidentalis var. pubescens Raf.
Fulton, Hempstead Co.; Hot Springs, Garland Co.
Galium. tinctorium L. 6083 Corning, Clay Co.
Houstonia lanceolata (Poir.) Britton. Lee Co.
(C. Woolsey). |
Caprifoliaceae:
Lonicera japonica Thunb. Japanese Honeysuckle.
Escaped in all parts of state and often becoming
troublesome. Naturalized from Asia.
Sambucus canadensis var. submollis Rehder. 22295
MeNab, Hempstead Co.
Viburnum affine Bush. Ozark region, on rocky cliffs.
Dipsacaceae:
Dipsacus sylvestris Huds. Teasel. Washington and
Benton Counties. Introduced in waste places.
140 Trans. Acad. Sci. of St. Louis
Cucurbitaceae:
Cucurbita foeditissima H. B. K. (Pepo foetidts-
sima Britton) Washington and Benton Counties.
Echinocystis lobata Michx. Micrampelis lobata
Greene) Washington Co.
Lobeliaceae :
Lobelia spicata var. hirtella (Ell.) Gray. 4407
Kureka Springs, Carroll Co.
Campanulaceae:
Sphenoclea zeylanica Gaertn. Lonoke, Lonoke
Co.; Stuttgart, Arkansas Co. An introduced annual
weed of rice fields.
Compositae:
Artemisia annua L. 4732 Cotter, Baxter Co.
Aster lateriflorus (L.) Britton. 26670 Helena,
Phillips Co.
Aster ericoides L. 4510 Eureka Springs, Carroll Co.
Aster ericoides var. pilosus (Willd.) Porter. 6812
Fulton, Hempstead Co.; 6883 Eureka Springs,
Carroll Co.
Aster salicifolius Lam. 8981 Fulton, Hempstead
Co.
Aster subsessilis Burgess. Benton County (type
locality, E. N. Plank).
Aster Tradescantu L. 4491 Eureka Springs,
Carroll Co.; Moark, Clay Co.
Bidens discoidea (T. & G.) Britton. 26733 Me-
Nab, Hempstead Co.
Supplement to Catalogue of Arkansas Plants 141
Brauneria pallida (Nutt.) Britton. 25022 Hazen,
Prairie Co.; Savoy, Washington Co. Common in
Ozark region.
Cacalia similis (Small) new comb. (Mesadenia
similis Small.) Benton Co. (type locality, E. N.
Plank); Savoy, Washington Co.
Chrysopsis mariana (L.) Ell. Southeast Arkan-
sas (County Agric. Agent).
Chrysopsis microcephala Small. 6847 Columbus,
Washington Co.
Cirsium horridulum Michx. 24977 Fulton, Hemp-
stead Co.
Coreopsis lanceolata var. villosa Michx. (C.
crassifolia Ait.) 26889 Lawrence, Garland Co.;
27056 Berryville, Carroll Co.
Coreopsis pubescens Ell. Savoy, Greenland, Wash-
ington Co.; 4490 Beaver, 6350 Eureka Springs,
Carroll Co.; Harrison, Boone Co.
Eupatorium incarnatum Walt. 4766 Cotter, Marion
Co. ; 14626 Texarkana, Miller Co.
Erigeron quercifolius Lam. Screeton, Prairie Co.
(Roland M. Harper).
Gaillardia lutea Greene. 10596 Washington, Hemp-
stead Co.
Helenium campestre Small. Little Rock, Pulaski
Co. (type locality, coll. by Hasse, 1885); Conway,
Faulkner Co. (H. E. Wheeler).
Helianthus divaricatus L. 12631 Mena, Polk Co.
26444 Magazine Mountain, Logan Co.; 29445 MeNab,
Hempstead Co.
142
Trans, Acad. Sci. of St. Louis
Helianthus heterophyllus Nutt. 6979 Heber Springs,
Cleburne Co.
Hymenopappus carolinensis (Lam.) Porter. 27052
Berryville, Carroll Co.
Lactuca canadensis L. Fayetteville, Washington Co.
Lactuca scariosa var. integrata Gren. & Godr.
Common over entire state.
Liatris squarrosa var. intermedia (DC.) T. & G.
23061 Hot Springs, Garland Co.
Matricaria suaveolens (Pursh.) Buchenau. Fay-
etteville, Wheeler, Washington Co. Introduced.
Mikania scandens (L.) Willd. St. Francis River
Swamps (S. M. Coulter).
Parthenium hispidum Raf. (Parthenium repens
Eggert). 5591 Beaver, 27059 Berryville, Carroll
Co.
Pluchea petiolata Cass. 314 Texarkana, Miller
Co.; 4756 Cotter, Marion Co. ;
Verbesina virginica Iu. 274 Rich Mountain (at
summit), Polk Co.
Lepachys columnaris (Sims) T. & G. Prairie
Grove, Washington Co.; 8024 Fulton, Hempstead
Co.; 8216 Benton, Saline Co.
Rudbeckia Brittonii Small. Fayetteville, Wash-
ington Co.
Senecio obovatus var. rotundus Britton. 7201
McNab, Hempstead Co.; 26866 Hot Springs, Gar-
land Co.; 24444 Little Rock, Pulaski Co.
Supplement to Catalogue of Arkansas Plants 143
Senecio plattensis Nutt. Prairie, near Hazen,
Prairie Co. (fide J. M. Greenman). A noxious weed.
Silphium Gatesii Small. 8048 McNab, Hemp-
stead Co.
Solidago amplevicaulis T. & GQ. 434 MeNab,
Hempstead Co.
Solidago caesia var. axillaris ae! Gray 6930
Jasper, Newton Co.
Solidago celtidifolia Small. 8461 Gifford, Hot
Springs Co.
Solidago arguta Ait. 26430 Magazine Mountain,
Logan Co.
Solidago hispida Muhl. 6912 Harrison, Boone
Co.; 26464 Lonsdale, Garland Co.
Solidago juncea Ait. 547 McNab, Hempstead
Co.; 4401, 4368 Eureka Springs, Carroll Co.
Solidago latifolia L. (Solidago flexicaulis L.)
4554 Hureka Springs, Carroll Co.; 4779 Cotter,
Marion Co.
Solidago Lindheimeriana Scheele. 20482 Eureka
Springs, Carroll Co.
Solidago pendula Small. Benton Co. (Type locality,
collected by E. N. Plank.)
Spilanthes americana var. repens (Walt.) A. H.
Moore. (Spilanthes repens. Michx.) 6085 Corning,
Clay Co.
Sonchus asper (L.) Hill. Lonoke and Arkansas
Counties in rice fields; Conway, Faulkner Co.; Fay-
etteville, Washington Co.
Trans. Acad, Sci. of St. Louis
Sonchus oleraceus L. Fayetteville, Washington
County and elsewhere (a common weed).
Tragopogon porrifolius L. Fayetteville, Wash-
ington Co. (as a weed).
Taraxacum erythrospermum Andrz. Conway, Faulk-
ner Co.
Vernonia missurica Raf. 300 Texarkana, Miller Co.
Xanthium glabratum (DC.) Britton. 316 Tex-
arkana, Miller Co.
Trans. Acap. Sci. oF St. Louis, Vor. XXV Puate VI
Supplement to Catalogue of Arkansas Plants 145
EXPLANATION OF PLATE
Puiate VI
Upper: Scouring Rushes (Equisetum suppers along railway em-
ankment near Palarm, Pulaski Co. . E. Wheeler.)
Lower: Arrow Leaf a latifolia) border of pond near Con-
way, Faulkner
146 Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
Pate VII
Upper left: Short Leaf Pine (Pinus echinata) in characteristic stand,
about 25 years old. Butterfield, Ark.
Upper right: Same growing on quartzite hills along Ouachita River,
Garland Co.
Lower eco Fringe Tree or Old Man’s Beard (Chionanthus virginica)
of Magazine Mountain, Logan Co.
Lower taki, Southern Buckthorn ere lycoides), Crowley's
Ridge, near Helena, Phillips Co.
Trans, Acap. Sci. or St. Louis, Vou. XXV PLate VII
inital.
Trans. Acapb. Sci. oF Str. Louis, Vout. XXV Puate VIII
Supplement to Catalogue of Arkansas Plants 147
EXPLANATION OF PLATE
Prate VIII
Left: Texan Saxifrage hi Saxifraga texana) probably the earliest spring
flow % wer, appearing late in February, Conway, Faulkner Co,
x
Middle: A very rare Wake Robin (Trillium pusillum) appearing in
il in oak woods on flinty soil. Tontitown, Washington Co.
Right: False Aloe (Agave virginica) flowering in June, near Fayette-
ville, Washington Co.
148 Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
Puiate 1X
Upper left: Indian Plantain (Cacalia tuberosa) near Imboden.
Upper right: ad Poison (Amianthium muscaetoxicum) near Imboden,
Lawre o. (H. E. Wheeler.)
Lower opie ae hay of the Ozark Chinquapin (Castanea ozarkensis)
ar F ington, Washington Co.
Lower ree Bark of Ozark Chinquapin, same tree as above.
TRANS. AcabD. Sci. oF Sr. Louis, VoL. XXV PLaTe IX
Trans, AcaD. Sci. oF St. Louis, Vor. XXV Piate X
Supplement to Catalogue of Arkansas Plants 149
EXPLANATION OF PLATE
PuatTe X
Upper left: Blue-fruited Hawthorn (Crataegus brachyacantha) near
Texarkana, Miller Co.
Upper right: Bark of same.
Below: Field of Spider Lilies (Hymenocallis occidentalis and Iris) ina
marsh near Palarm, Pulaski Co. (H. E. Wheeler.)
150 Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
PiatTe XI
Upper left: Honey Locust (Gleditsia triacanthos) near Helena, Phil-
lips Co.
Upper right: Bark of same.
Lower grin White Ash (Fraxinus americana) growing from opening
trunk of Satiow Post Oak (Quercus encter Fulton, Hemp-
Co,
Lower right: Azalea (Rhododendron canescens) in April near Hot
Springs.
Pirate XI
ry eee, 4
uls, Vi
I
s. Acapb. Sci. oF Sr.
TRAN
=
‘yy
Puate XII
XXV
OL.
rT
Sci. oF St. Louis, \
TRANS. ACAD.
Supplement to Catalogue of Arkansas Plants 151
EXPLANATION OF PLATE
Priate XII
Upper left: Blazing Star (Liatris elegans) and Button Snakeroot
Eryngium yuccaefolium) in a meadow near Palarm, Pulaski
Upper right: Chicasaw Plum igeiaas angustifolia) growing on loess
s, Helena, Phillips
Lower ets ogi Leaf Pine Feet echinata) West Mountain, Hot
Lower right: French Mulberry ( Callicarpa americana) a shrub with
br -epubers lavender colored berries in autumn. Helena, Phillips
152 Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
PuatTe XIII
Cone Flower (Brauneria pallida) in a meadow in May,
shington Co.
Upper right: Red Buckeye Shin discolor var. mollis) flowering in
May, Central Arkans
Lower left: Prairie a 8 ee purpureum) in June, North-
west Arkansa
Upper left:
Wa
Lower right: Rough ee [Mecente scabra) Arkansas border and
Southwestern Missouri in
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Supplement to Catalogue of Arkansas Plants 153
BIBLIOGRAPHY ON FLORA OF ARKANSAS.
(Chronological)
Bradbury, John. Travels in the interfor of North America in
the years ae _ 1811. Liverpool, 1817. (Mentions
Several plan om “Upper Louisiana” and Great
Prairie, probably ria Arkansas, seen on trip down Mis-
sissippi Rive
- Schoolcraft, Henry R. Journal of a tour into the interior of
telat and Arkansas. (1818-1819). London (Phillips).
Nuttall, Thos. Journal of travels into Arkansas territory dur-
ing mo year 1819, with tigger observations on the
manners of the aborigines . 236. Phila. 1821.
rnin cae of some new species of plants recently
introduced into the gardens of Philadelphia from the
Arkansas territory. Journal Philadelphia Academy o
Science. 2:114-23; Phila. 1821. (Includes description of
twelve species of Arkansas ane Ss.)
~————— Description of two new genera of the natural order
Cruciferae. Journal Philadelphia Academy of Science,
5:132-35. 1825. (Described: Selenia aurea Nutt. and
Strepthanthus maculatus Nutt.) Plates 6-7.
Featherstonehaugh, G. W. Geological report of an examination
made in 1834 of the elevated country sorcenn the MiIs-
rie and Red Rivers. Exx-Doc. No. 151, 23d Cong, 2nd
Vol. IV. 97 pages. Washington (Gates and
35.
- Nuttall, Thos. Raiegeconsy towards a of the territory of
Arkansas, Trans. Am. Phil. eg oan Series) 2:139-202.
Phila. "1887
- Lesquereux, Leo. Botanical and paleontological report on the
geological state survey of Ar as. Rep. Second Geol.
in of Ark. pp. 295-399. Phila. 1860.
Butler, Geo. D. Additions to the flora of Arkansas. Bot. Gaz.
2:104. 1877.
Harvey, F. L. Asplenium Bradleyi. Bot. Gaz. 5:15, 1880.
Some Arkansas ferns. Bot. Gaz. 5:39. 1880.
Early plants. Bot. Gaz. 5:56. 1880.
(Coulter, J. M.) Southwestern plants. Bot. Gaz. 5:84.
Harvey, F. L. Notes from Arkansas. Bot. Gaz. 5:91-3, 1880.
Distribution of Nymphaceae in Arkansas. Bot. Gaz.
: 80.
1880.
Warder, John A. Notes from Arkansas, Bot, Gaz. 6:188. 1880.
Harvey, F. L. Ferns of Arkansas. Bot. Gaz. 6:189-90. 1880.
Ferns of Arkansas. Bot. Gaz. 6:213-15. 1880.
—————— Some Arkansas Trees. Bot. Gaz. 6:215. 1880.
oo
or
ow
~1
i)
oo
. Williams, J. te
Trans. Acad. Sci. of St. Louis
Leavenworthia in S. W. Missouri and N. W. Arkansas.
. Gaz. 6:230. 1880.
The flora of Arkansas. Am. Nat, 15:388-89. 1881.
Forest Notes. Bot. Gaz. 6:273. 1881.
A second Spring in Arkansas. Bot. Gaz. 7:12. 1882.
Selenia aurea. Bot. Gaz. 7:57. 1882.
Forestry Notes. Bot. Gaz, 8:355-56. 1883.
Carya myristicaeformis, Bot. Gaz. 9:195. 1884.
Dioclea Boykinii. Bot. Gaz. 9:196. 1884.
Notes on forest trees. Bot. Gaz. 10:279-80. 1885.
Some abnormal Rudbeckias. Bot, Gaz. 10:296. 1885.
. Sargent, . S. Report on the forests of North America Sees rth
th
Mexico). 10th U. S. Census (Arkansas) 9:543-4
Washingt. 1884.
Map of Arkansas rene the distribution of the forests.
10th U. S. Censu
. Harvey, & L. Arboreal flora of Arkansas. — Jour, Forestry.
1:413-24 and 451-58. June-July.
. Call, R. Ellsworth. Notes on the native forest trees of eastern
Arka
nsas. Proc. Ia, Acad. Sci. 76-85. 1887-89.
(Branner, J. C.) Flora of Arkansas with additions by W. a
Blatchley, 1889, and index; also notes of others added a
various ti
Manuscript in University of ” arkansas
library. (No date.
. Branner, J. C. and F. V. Coville. A list of the plants of Arkansas.
18
Annual Report for 88. Arkansas Geol. Survey,
4:155-242. Little Rock. 1891.
. Coville, F. V. Notes on the Botany of Arkansas. Annua
al
port for 1888 Arkansas Geol. Survey, 4:243-252. ens
Rock.
Call, R. sg irdhe ac Notes on the forest trees of the Cromer?
Ridge region. Annual Report for 1889 Arkansas Geo
Survey, 2:183-202. Little Rock.
Distribution of timber and its relation to
underlying =P formations. (Magnet Cove — 2
kaunas Report for 1889, Part 2. (Igneous Rocks
kansas Geo Garris for 1890. 2:167-170. Little ee
Harvey, F, Bat Varieties of Ranunculus abortivus L., Bull. Torrey
. Club. 19:93-94. March, 1892.
Bush, B. F. Bot of some southern swamps. Garden and
Forest 10: 51416, Dec. 1897.
Plank, E. N. Concerning the plants of southwestern Arkansas.
Plant World. 2:45-47. 1898-99. :
Olmstead, F. E. A I plan for the forest lands near Pine
Bluff,
U. S. Bureau of Forestry Bull. 32.
Washiantan. 1902.
we
wo
- Ashe, W. W. Further notes on woody plants.
1925.
Supplement to Catalogue of Arkansas Plants 155
- Coulter, S. M. Ecological comparison of some typical swamp
rea. Fifteenth Annual Report Missouri Botanical Gar-
den 39-71. 1904.
- Record, Samuel J. The forests of Arkansas. For. Quart. 5:296-
301. S. 1907.
The forest resources of Arkansas. Mo, Pac, Land Office,
Little Rock. 1910.
- Howell, A. H. Birds of Arkansas. Bull. No. 38, Biological Sur-
., Washington. 1911. (Contains
several botanical notes.)
- Harper, Roland M. Botanical evidence of the age of certain ox-
bow 6 1912
lakes, Science, N. S. 36:760-761. Nov. 29,
sae notes on the _ coastal saabae of
Arkansas. Plant World, 17:36-48. Feb. 1914.
Some undescribed prairies in northeastern Arkansas.
Plant World 20:58-61. Feb. 1917.
Sargent, C. S. Notes on North American trees. II. Carya. Bot.
Gaz. 66:229-258. Sept. 1918.
- Palmer, E. J. The forest flora of the Ozark region. Journal Arnold
1921.
rboretum 2:216-32. Apr.
he Red River forest at Fulton, Arkansas. Journal
avede Arboretum 4:8-33. Jan. 1923.
Ashe, W. W. Further notes on the trees and shrubs of south-
eastern United States. Bull. Torr. Bot. Club. 50:359-
363.
Notes on woody plants. Jour, Mitchell Soc. 50:43-48.
Aug. 1924.
- Palmer, E. J. The ligneous flora of Rich Mountain, Arkansas and
Oklahoma. Journal Arnold Arboretum 5:108-134. Apr.
1924.
Am, Fern Jour.
Two interesting ferns from Arkansas.
14:39-41. Apr.-Jun. 1924.
Buchholz, John T. Notes on Arkansas Pteridophytes.
Jour. 14:33-38. Apr.-Jun. 1924.
Am. Fern
- Shreve, ean W. Iris cristata. Flower Grower 11:301. Aug.
Flower Grower. 11:385-
Native irises of the Ozarks.
386. Oct. 1924
Buchholz, Rages T. and Wilbur R. Mattoon. Common forest trees
f Arkansas. Bull. Agr. Extension Division, University
(Little Rock Office.)
ae godine No. 180. Nov. 1924.
Torreya, 25:10-11.
Jan.-Feb.
a
3 2 j bs
e
Transactions of the Academy of Science of St. Louis
He
Volume XXV, No. 7 Se
THE ECOLOGY OF A SHELTERED CLAY
BANK; A STUDY IN INSECT
SOCIOLOGY
PHIL RAU
.
THE ECOLOGY OF A SHELTERED CLAY BANK;
A STUDY IN INSECT SOCIOLOGY.
Pum Rav.
A. Introduction.
B. Interrelations of animal life.
(a) Fauna of the unit.
(b) Pioneer life.
(c) Renters.
(d) Visitors.
(e) Parasites.
C. Relation of population to environment.
(a) Relation to temperature.
(b) Relation to light and sunshine.
(c) Relation to cold, cloudiness and darkness.
(d) Relation to rain.
(e) Relation to plant life.
(£) Death by violence and natural death.
(g) Relation to soil conditions.
D. Concluding remarks.
158 Trans, Acad. Sci. of St. Louis
(A) INTRODUCTION.
“‘Tiving things are real things—but their reality 1s
in their interrelations with the rest of nature, and not in
themselves.’’—Brooks.
The ecologist, when he wishes to study the interrela-
tions of a fauna to a restricted area, usually selects a
virgin plot, uncontaminated and undisturbed by the hand
of man. The area which I have studied is unique in that
it has been created by man. These pages will reveal that
the invertebrate population readily responded to this
man-made habitat and lost no opportunity to utilize il to
advantage. Despite the fact that the clay bank here re-
ferred to was artificially built, the attraction to it of the
life about the area for nesting and other purposes was
legitimate, and this made the place a biotic unit of un-
usual interest.
I have given to the unit a layman’s study of ecology,
made without the technical and refined instrumental ex-
aminations and prepared without the use of new and
complex terms so often introduced into a study of this
kind, which, in my opinion take commonplace phenomena
and place them beyond the understanding of the layman.
By trying to become familiar with complex and unusual
terms, one often loses the thread of the story of the in-
terrelations of the inhabitants. In other words, this
paper is, I hope, analogous to the study of the stars
through an opera glass, instead of a study by mathe-
matical formulae.
The ecological unit under discussion lay in a little
valley at the station of Wickes, twenty-two miles south
of St. Louis on the Iron Mountain Railroad. The Mera-
The Ecology of a Sheltered Clay Bank 159
mec River empties into the Mississippi about one-third
of a mile northeast of this point. This region is attrac-
tive to fishermen and as a result about a dozen club-
houses have been built near the station. Some of these
cottages were built upon posts or stilts, thus exposing
the earth beneath them. Two such houses were situated
on a northward slope, but there was no perceptible life
in the earth under them. This was probably due to their
shaded condition and the friable nature of the earth
there. In the construction of another of these houses, a
quantity of clay had been excavated from the cellar and
had been shoveled under the porch to save the expense of
hauling it away. In the twelve years that this heap of
subsoil had lain there, it had become packed to a very
hard consistency. The house faced the east where it
received the benefits of the morning sun. The porch was
four to five feet above the ground, and thus admitted a
flood of light in the morning and midday, but excluded all
the afternoon sun, and afforded protection from the
weather. This bank of hard-packed yellow clay was
about eighteen feet long, and varied from thirty to thir-
ty-six inches in height. The wood-boring inhabitants of
the porch, the life in the clay bank, the insects among the
rubbish on top, the occupants of the rambler-rose stems in
front of the bank, and the occasional visitors all consti-
tuted a legitimate biotic unit, the subject of this study.
(See Fig, ie
In May, 1917, when the unit was first discovered, it was
noted that the northern half of the clay bank was riddled
With holes, while the south half was not so, but at the
extreme south end a few turrets of a wild bee were found.
Figs. 3 and 4 show the south and the north half of
the clay bank respectively, and the enormous amount of
160 Trans. Acad. Sci. of St. Louis
activity which occurred in the north half in contrast to
that in the south half will be seen by the contrast of the
great number of holes in the one to the lack of them in
the other. Before the summer was far advanced, the
cause for the occupancy of one portion of the bank and
not of its opposite half was clearly apparent. ‘A glance
at Fig. 2 will show how the central and south rose-
bushes had been trained to overshadow the southern por-
tion of the clay bank, while the third bush had been so
bent as to leave the northern half exposed to the eastern
sun.* This shows beautifully the relation of sunlight
to the activities of Hymenoptera. The loose appellation
usually applied to them, ‘‘Children of the Sun,’’ is here
seen to have truthful significance. It is generally known
that bees and wasps love the sunlight, but a glance at
Figs. 3 and 4 will forcibly emphasize this impression.
While this clay bank was small and unimposing, its
riddled portions indicated that many bees and in all
probability their parasites were potentially present, and
for an ecological unit for intensive study nothing better
could be wanted. Accordingly, frequent visits were
made to the spot from June 5 to October 3, 1917, and
several trips were made there for comparative data dur-
ing the summers from 1918 to 1921.
When one looks at Fig. 2 and thinks of the life, love
and death tragedies enacted within the limits of an area
of a few square feet, one can see very clearly a literal
example of the statement by Adams: ‘‘ We may profitably
compare an association of animals in a given habitat to
a play upon the stage. The environment corresponds to
*The photograph for figure 2 was taken in October, when most
of the leaves had fallen, and so does not give an adequate idea of
the density of the shade.
The Ecology of a Sheltered Clay Bank 161
the stage, the dominant members of the association cor-
respond to the leading characters; the secondary species,
always present, to the essential but subordinate charac-
ters. The individual animals adjust themselves to one
another, especially to the dominant forms, and to the
environment, as the personalities in the play adjust them-
selves to the dominant characters, to one another, and to
the environment.. In both groups some individuals are
dominant, some used and useful, some tolerated, others
pick up the crumbs, still others are predatory or parasitic,
and all must be mutually adjusted to one another and to
the environment.’’”*
How this classification of various types fits into what
we here record, the reader may presently see for himself.
(B) INTERRELATIONS OF ANIMAL LIFE.
(a) Fauna of the unit.
If all the bank’s a stage, and all the six and eight-
legged creatures merely players, it is fitting that we
should give now a cast of characters, in the order of their
importance, and follow the careers of the dominant ones
for the five years, 1917 to 1921. Early in the work it was
seen that the many insects did not use the clay bank and
the environs in the same way, but they were easily classi-
fiable into four distinct groups.
Group 1. Pioneers. The permanent dwellers in the
clay bank and environs, in a general way the pioneers.
Group 2. Renters. The insects less hardy and more
ease-desiring than the pioneers; those which rented
or appropriated the abandoned dwellings of the pioneers.
They might be called squatters.
*Guide to the Study of Ecology, p. 47. 1913.
=
162 Trans. Acad. Sci. of St. Louis
Group 3. Visitors. The insects and animals which
dropped into the community accidentally, or in quest of
shelter or food. They very often influenced the inhabi-
tants, the members of groups 1 and 2, in two ways: by
eating them or by becoming food for them.
Group 4. Parasites. This contains the names of
parasites whose hosts are listed in groups 1 and 2.
(b) Pioneer life on the clay bank.
The pioneers, those which blazed the trail, those which
first came from elsewhere and discovered the clay bank
and homesteaded it, and made it easy for others follow-
ing to live near them or in their old abodes, or to para-
sitize their children—these are first considered, and are
listed in order of their importance. With this pioneer
life as a foundation for a community, the other chapters
that follow will point out their interrelations, sometimes
simple, sometimes indifferent and sometimes complex.
The carpenter-bee, Xylocopa virginica. Drury.”
In the Spring of 1917, the place was visited at in-
tervals to fix the date of the appearance of the first life.
The carpenter-bee, Xylocopa virginica, was the first to
appear. On June 15, perhaps a dozen of the insects
were at work enlarging the tunnels in the wooden rafters
of the porch above the clay bank. They were all doing
precisely the same thing at this time, enlarging the holes
and kicking out the sawdust. The flickering streams of
golden grains falling in the sunlight from sources un-
seen would have attracted the attention even of persons
unaccustomed to observations of this kind. Judging from
*All of the material has been identified by expert entomologists,
ose names appear in brackets along with the specific name of the
i;
The Ecology of a Sheltered Clay Bank 163
the piles of sawdust here and there on the ground, this
work must have been going on for a good many days.
Hence the emergence of the bees was simultaneous, but
whether from hibernation or from the pupal stage I do
not know, since I could not ascertain just how this species
spends the winter, unless I used an ax on the rafters,
and this I had not the immoral courage to do. However,
it was plain from the first that no new tunnels were being
made, but that old ones were being enlarged.
The activities of the bees in tunneling and pollen-gath-
ering increased, although not their number, up to July
16, when they almost completely disappeared. This was
probably the end of the first generation of 1917, regard-
less of whether they had hibernated or had emerged as
new adults in the spring.
On July 30, other adults appeared—probably the first
of the second generation. On this first day, two females
were at work; on the next day four were in evidence,
and the numbers continued to increase during the next
few days, until by August 20 they were more abundant
and seemed more industrious than the population earlier
in the season. They were so intensely busy cutting out
the tunnels, and they threw down the yellow dust to the
ground in such abundance that the pits of the ant-lion
larvae beneath suffered complete obliteration. They
were seen in the same activity and numbers up to Sep-
tember 7 (with certain pauses in their work due to mete-
orological conditions described elsewhere). The three
nights before September 12 were very cold; the insects
about the unit were dwindling in numbers, and those
which still lived were slow to come out of their burrows.
That day it was 2 o0’clock in the afternoon before the first
carpenter bee made its appearance, but her sisters re-
164 Trans. Acad. Sci. of St. Louis
mained within, in either temporary or final rest. Again
on September 23 only one carpenter-bee was seen. At
this date all the other life was proportionately reduced,
as was to be expected amid unfavorable weather condi-
tions. A visit to the spot ten days later, October 3, re-
vealed several dead specimens of these bees and other
species about the bank, but no living ones. In all prob-
ability they were either dead in their burrows or closely
huddled together, ready to hibernate for the winter.
In the year 1918, as I shall relate elsewhere, all the
more important species in the unit appeared earlier than
in the previous year. In 1917, the first carpenter-bees
were sawing their tunnels on June 15; in 1918, on May
28 not only were they at work, but they had already com-
pleted their burrowing and were busily gathering the
yellow pollen from the wild roses growing some distance
away. The big shining black bee, heavily laden with
bright yellow pollen, made a conspicious bit of color, and
was as pretty as the blossom itself. An examination
showed that no new tunnels had been made, but the old
ones from the last year had been again used.
My next visit on June 28 found no carpenter-bees out;
this was probably the period between the two genera-
tions as was July 30 in 1917, when the first generation
was late by a corresponding margin of time in getting a
start. On July 17, when it seemed time to expect the
second generation, I visited the colony, and found one
lone carpenter-bee at work; July 31 likewise found only
one bee out.
All of the evidence so far in 191% indicated a marked
reduction in the population of carpenter-bees. An abun-
dance of shedding-skins of the silver-winged parasite,
Argyromoeba tigrina, adhered to the wooden rafters
The Ecology of a Sheltered Clay Bank 165
about the bees’ tunnels; hence it is probable that this
parasite was largely responsible for the small number
of bees, as well as of the mud wasps, Monobia quadri-
dens, and the grass-carrier wasps, Chlorion (Isodontia)
auripes, which occupied the old tunnels of the carpenter-
bees. The details given later under the subject of para-
sites will show an enormous increase at that time in the
number of adults of the A. tigrina parasite,
So the record for the second generation continued dis-
mal; on August 31, barely two bees were busy, and by
the middle of September none at all were to be seen.
During the summer of 1919 no observations could be
made, so we do not know by what struggles or turns of
fortune Xylocopa again came into her own, but in the
following spring, on May 27,* 1920, they were once more
fairly abundant. Their emergence had probably just
occurred when I arrived upon the scene, for they were
not tunneling, neither were they carrying pollen, but
both sexes were in a gleeful mood of courtship.
Several days later others were often seen refresh-
ing themselves on the blackberry blossoms, while sev-
eral returning females were so heavily laden with green
pollen that much of it would spill in a delicate shower
as they tried to gain entrance to the home. During the
week of June 13 to June 19, eight mothers were at work
on their nests. Since only these eight remained to work,
out of the large number in the courtship dance, it leads
one to suspect that a very large proportion of the latter
group were males. During the days of June 28th and
29th, I kept a very careful watch and was startled by the
revelation which met my eyes; during the two hours
*In 1922 they were flying about as early as April 7. One newly
dead bee was taken from a spider’s web.
166 Trans. Acad. Sci. of St. Louis
from five to seven p. m., 35 adult Xylocopa returned to
the colony and quietly crept into their burrows! This
certainly shows how an observation must be made
from every possible angle before one may dare to be
satisfied with an answer. Fourteen of these were cap-
tured and examined; twelve were females and two were
males. The presence of the males made me wonder
whether they were the remainder of the first generation,
or the forerunners of the second. I cannot imagine just
what was their business abroad which kept them away
from home all day thus, day after day, at a time when
they were not gathering provisions of any kind for the
nest. May it be that they were out seeking food to gorge
and fatten themselves before beginning the arduous
tasks of nidification? From this time on the population
again waned, until by the middle of September they were
seldom seen. Long after the others had disappeared
from view, one lone mother was seen going into her
burrow on October 13.
Just as the mining bees make way for numerous other
species by bequeathing to them their old burrows, 8o
Xylocopa makes it possible for two species of wasps,
Chlorion (Isodontia) auripes and Monobia quadridens
to inhabit the locality. For details of the behavior of
these sub-tenants and for details of parasitism on the
carpenter-bee, and for the relation of the bee to weather
conditions, see later pages.
The mining bee, Anthophora abrupta Say [S. A.
Rohwer].
The most important of the pioneers getting a foothold
in the clay bank were two species of mining bees, one
supplementing the other beautifully in the point of time.
The Ecology of a Sheltered Clay Bank 167
The first to appear each year was Anthophora abrupta,
and when its life cycle had run its course before the
middle of the summer, the white-banded bee, Entechnia
taurea, made its appearance. Other species of bees ap-
peared from time to time, but none reached the impor-
tance that these two species attained.*
On June 25, 1917, it was found that several specimens
of the mining-bee, Anthophora abrupta (Fig. 5), had
emerged from their winter sleep in the depths of the
clay bank, and were busily coming in and going out of
their burrows. Some of these activities were directed
to burrows which had mud chimneys at the openings, and
some to others which had none. Some of these bees came
in heavily laden with pollen; others had their gullets full
of water; others were kicking out loose moist earth from
their burrows and letting it fall to the ground below.
Some were working within the burrows, and at intervals
would come backing out with a ball of soft mud under the
chin, toss it back to the hind pair of legs, and with only
these appendages would fashion the wonderful little
chimney which crowned the entrance; others were fre-
quenting the puddle of water in a wagon rut near by,
getting the water with which to moisten the hard yellow
clay. Quite probably all these bees had emerged within
a few days prior to June 25, and they were already at
work in various stages of nest-building or provisioning.
June 28 was my next visit, and this was after a heavy
Shower. There was little activity among the bees; most
of them were waiting indoors until the sunshine should
warm up the bank, but several had learned to make a
*In 1919, a third and new species of mining bee, Anthophora
a oi ractically the same life cycle, and ap-
peers at the’ odes S cant a few days later than
a.
168 Trans. Acad. Sci. of St. Louis
lucky turn out of this misfortune; instead of going some
distance to the roadway puddle for their water, a dozen
bees with their long tongues protruding were lapping
up the drops of water from the vegetation nearby.
Frequent visits showed the activity of the A. abrupta
bees to be the same without change in numbers up to
July 12. Thus the entire population lived practically
simultaneously, without individual variation in time.
During the four days from July 12 to 16, the number of
workers dwindled alarmingly from day to day. This sud-
den dropping off led me to believe that I was witnessing
the death and the end of the first generation. I hoped
that the second generation would come soon enough and
be large enough to give me material for certain homing
studies without the danger of exterminating them. The
next few days saw the demise of the last few survivors;
but my anticipation of a second generation was in error,
for no others appeared that year. Furthermore, in the
years following I found only one generation of these
bees each year, and that with a remarkably short life
eycle. In this 1917 brood, the duration of adult life was
less than thirty days, the first bees emerging a few days
before June 25, and the last ones dying about July 16.
While daily visits during the following week proved
that A. abrupta had entirely disappeared, these daily
visits also made it conspicuously clear that another
species of turret-building mining bee, Entechnia taurea,
had emerged from the bank and was busily engaged in
nest building activities. And in precisely the same way
in the following years it was observed that just as soon
as A. abrupta had finished her labors and gone the way
of all flesh, then the white-banded bees appeared upon
the scene. The interrelations of the two species of bees
The Ecology of a Sheltered Clay Bank 169
in point of numbers from year to year show very fine
adjustment. As one species increased progressively from
year to year, the other decreased. The details of this
phenomenon are given on later pages.
While 4. abrupta appeared about June 25 in 1917, a
visit on May 28, 1918, showed this species already out
and active. Early emergence this year affected most
Species inhabiting the bank. Later I shall discuss the
relation of this emergence to meteorological conditions.
But even at this early date, the A. abrupta bees must
have emerged five or six days previously because the
work on their chimneys was in many cases far advanced
or completed. Having been born a month earlier in the
year than their parents of the preceding year had been,
the population had the same number of days to their
adult life and lived, builded, and many had died before
June 28, Then, within three days after their disappear-
ance came the white-banded bees again, following with
all the precision of clock-work! The interesting item
was that Anthophora abrupta had increased in numbers
from 22 nest-building mothers in 1917 to 92 in 1918.*
Not all of the 92 mothers had nests with turrets; only
52 had full-sized chimneys, 15 with half-sized ones and
25 nested in burrows devoid of chimneys. This increase
was, as we shall later show, very decidedly at the ex-
pense of the white-banded bees whose numbers this year
rapidly decreased. What should we expect in the fol-
lowing year, and what was the cause of the change in
dominance?
*It , that any accretions in numbers from
ene tu this cis We A. abrupta were progeny from the
t
Since the latter species diminished in the bank from year to year,
is evident that no new stock took up their abode there.
170 Trans. Acad. Sci. of St. Louis
In 1919, circumstances were such that I could pay my
first visit to the clay bank only on September 6. There-
fore, I could not get their date of emergence, bunt they
had left for me a clearly-written record of their abun-
dance on the face of the bank. I counted 232 turrets,”
which indicated that at least that number of mothers had
been at work. Since a small portion of the mothers used
the burrows without turrets, and since some may have
been broken down, this is a conservative number. I am
sure that none were from the previous year, for I had
harvested the entire lot in the autumn for my cabinet
and study. A recapitulation of the figures is of interest:
in 1917, 22; 1918, 92; 1919, 232.
In 1920, the first visit to the clay bank was made on
May 28. Until 9:45 a. m. no bees appeared; then the
sun warmed the bank and I saw two A. abrupta flying
before the bank and entering numerous holes, one after
the other. During the next fifteen minutes I sat before
the bank and saw three others emerge from the burrows
and dart away on the wing. The white faces of all five
proved that they were males. A shrill voice within a
burrow caused be to rivet my attention to a certain spot,
where soon a sixth white face made its appearance. So
here I was at hand for the first time to see the males
emerge! There were no females about. There occurs
in this species that which we see in so many species of
insects—the priority of the emergence of the males.
About noon the first female was seen going into burrow
after burrow, and a little later a second one was doing
precisely the same, while the males began to go away to
*This had been an especially wet year and the bees could get
water for digging. In dry years, as in 1922, hardly any of them
made turrets.
The Ecology of a Sheltered Clay Bank 171
the blackberry blossoms, or to rest listlessly upon the
nearby vegetation. The females moved about with an
air of hesitation, of uncertainty. All this showed that
the emergence of this species had probably begun on
that day, so, judging by the last year’s population, we
had reason to expect that the next two days would show
an exodus of great magnitude. Unfortunately, I could
not be present during the next few days.
On June 13, a very different type of activity was ap-
parent. Hundreds of turrets were already completed
and hundreds of busy mothers were flying in and out of
their nests. In two weeks wonders had occurred. The
males had come and gone—not one was to be seen.
Judging by the progress of the work each activity must
have been practically simultaneous for the whole group.
Thus, all nests were in the same stage of construction,
indicating that the work had begun simultaneously;
hence probably emergence and mating had occurred
for all within a day or two.
The amount of work which they had already done
was enormous, such as could only have been accom-
plished by their toiling as they do from sun to sun. At
5:30 a. m., one day I saw some of them beginning their
work, and at 7:30 p. m., when it was almost too dark in
the shadows to find their burrows, they came home and
crept in. Later in the season they were not so in-
dustrious early and late on cool or cloudy days. : It is
their habit to carry water in the gullet, with which to
reduce the hard yellow clay to workable mud. When one
takes a bee in a test-tube, it disgorges the water on the
glass in its futile fury. On one occasion, several bees
in a test-tube disgorged so much water that they became
172 Trans. Acad. Sci. of St. Louis
quite drenched and made their elegant pubescence all
sodden.
The notable feature was that this species had again
increased enormously. Their distribution over the face
of the clay bank was interesting: at the extreme south
end were 10 turrets, in the central part, 275, and at the
extreme north end, 390, making a total of 675 in three
rather distinct groups. Besides this, more than a hun-
dred bees had nests with no turrets and during the week
I saw about 75 more chimneys constructed. This may
indicate that each mother may make more than one nest
in a season.* Many of the bees were bringing in pollen,
others were building, and yet many more than three-
fourths were at this time coming in apparently empty-
handed. I think that they moisten the pollen with nec-
tar, and this would mean many trips which to an ob-
server would appear empty. Then again each cell is
varnished inside to a certain depth which gives it a
smooth glossy finish. This is no doubt for the purpose
of making the brood cup impervious to water; at least
when one drops water on the inside from a pipette,
the drops retain their shape, and if the entire cup is
filled, it retains the water for a long time. On the con-
trary, if a drop be placed on the outside of the cell, it
immediately spreads and makes mud out of the mass.
This varnish is probably made from plant resin that
is carried at the time when they seem to be coming home
empty handed. This bee, and the two other species of
mining bee, do more than merely make a hole in the
. ground and fill one end with food; they actually make
mud honey-cells which ean be taken from the burrow in-
*There may be dull seasons, e. g. cold or dry, like sacs when it
would “eg impossible for a female to make more than on
The Ecology of a Sheltered Clay Bank 173
tact (Fig. 11), and all three species varnish the interior
with the waterproof substance already referred to.
As noted above, they customarily work from early
till late. On June 17, 1918, when the temperature*
dropped from 98 degrees to 79 degrees Fahrenheit, their
activities were greatly lessened. Less than a dozen as-
sumed activity at all.
I tried to find where they got their pollen. On a per-
simmon tree some two hundred yards away, with its
blossoms already deteriorating I found many bees, in-
cluding this species, getting nectar. The two popula-
tions, bees and blossoms, waned at the same time. In
1921, notes on the relation of this species to the persim-
mon blossoms were made, and details are given else-
where. What other sources of food supply they may
have had I did not learn.
From May 28 to June 13 of the year 1920, they had
built turrets to the enormous number of 675, a substan-
tial increase over the number for the previous year. On
July 21, only a week later, these short lived creatures
were almost gone; only two live females could be found,
the remnant of the recently noisy and mighty throng,
and dead specimens of A. abrupta were scattered all
about. In 1921, Man was again a factor in regulating
the abundance of Anthophora life; the tenants of the
“‘Ham and Bud’? club-house unwittingly caused the death
of hundreds of these little creatures, by merely going
away and leaving the door of the screened porch open.
The bees, in their circuitous flights, entered this open
door at the head of the stairway (Fig. 8, point x), and
Were not able to find it again to make their way out; thus
the screening which was intended to keep insects out,
ee
*U. S. Meteorological Summary for June, 1918.
174 Trans. Acad. Sci, of St. Louis
kept these in. Within the porch was a pitiful sight; in
the two sunniest corners of the porch was literally a
drift of dead bees. In their desperate attempts to es-
eape, they had directed all their efforts against these two
points of maximum sunlight, and even while I watched,
at the southeast corner in the bright sun were one hun-
dred and fifty mothers trying frantically to escape. It
was with a feeling of satisfaction for the privilege of
doing reparation for the wrongs done by my kind that
I picked these up one by one, and liberated them.
It seems strange, and yet it is logical, that these crea-
tures could not find their way out. In the first place they
had entered by chance, instead of direct quest, which
would entail flight of orientation and the formation of
memory images of the open door. Second, these are
sun-loving creatures, following the greatest intensity of
light, so it is easy to see how this factor would guide
them in their efforts to escape. This failing is common
not alone to Anthophora abrupta, but to other insects as
well; several specimens each of Arotes amoenus, Cress.
(R. A. Cushman), Megarhyssa lunatrix, Fab. (R. A.
Cushman), and Tabanus lasiophthalmus, Macq., were
also found there. A humming-bird had been trapped in
just the same way, and its little carcass too lay in the
sunniest corner, with its delicate bill thrust through the
wiring, showing its frantic efforts to escape.
An actual count of these dead bees gave me 18 males
and 703 females. This, plus about 300 females then at
work, shows the enormous proportions to which this
population had grown. Furthermore there should be
added to this total some 60 specimens from this group
that had been lost in certain of my experiments. One
wonders what would have happened to the bank in the
The Ecology of a Sheltered Clay Bank 175
following year if all these mothers, over 1000, had been
left alive to propagate the proportionate population.
It seems hardly possible that the clay bank could have
held them (and we know that crowded housing condi-
tions lead to ill health and ill morals). Would they,
under such pressure, have disseminated over other areas?
At the end of the season the results of this catastrophe
could be seen, for on June 24, 1921, we could count only
175 nests, as compared with the 675 counted for June 13,
1920.
The white-banded bee was now present, but in shame-
fully reduced numbers. One can hardly say that the A.
abrupta in ever increasing numbers had simply crowded
out the white-banded bees, because there was so much
unusued space about the clay bank, and the two species
occurred at different times, so they could not have been
in competition for the same flowers. It was almost wholly
a question of parasites, and this is treated elsewhere.
Other details on the life history of this bee, and the re-
lation of the species to moisture, light, etc., are given in
later pages.*
White-banded Mining-bee, Entechina taurea Say [J.
C. Crawford].**
The preceding pages show how the mining-bees, An-
thophora abrupta, rose in five years from a colony of
few individuals to a great and important population.
We have also seen how J. abrupta emerges early 1n sum-
mer, does its life-work in about thirty days and is done
he: logical aspects of the life
Wess or ths ee othe Oe camerien species of bees and
i : however,
will be treated biologically in other papers to follow. It is,
often difficult to draw the line between the two aspects of the study.
**Mr. Rohwer writes that this bee is now known as (Entechnia)
Melitoma taurea Say.
176 Trans. Acad. Sci. of St. Louis
for that year. These pages will show that each year
immediately after this, the second mining-bee, Entechnia
taurea, comes upon the scene and continues the mining
activities. The bees were observed for five years, and
their numbers ran practically in inverse ratio to those
of A. abrupta; as the one increased, the other decreased.
These white-banded bees (Fig. 7) emerged from the
clay bank in 1917 on about July 16, and took up their
work on the spot. This species likewise made turrets
over its burrows, but they were a little smaller in size
and of finer texture than those of A. abrupta. Like A.
abrupta, these bees did not seal their burrows, either at
the end of the chimney or the mouth of the burrow.*
The males of this species do not seem to emerge before
the females; or at least, if they do, they do not die earlier,
for on July 311 saw a lot of frolicsome males nea
at the north end of the bank.
The activities of the white-banded bees ne
throughout the summer. They never swerved from their
self-restricted nesting areas, two spots at the extreme
ends of the clay bank. By August 14, the burrows of
these bees were becoming somewhat abundant, and the
dancing males had now spread themselves to the south
end, where they buzzed and danced about the busy fe-
males coming home heavily laden with pollen. While
some of the latter burrowed horizontally into the face of
the bank, others dug vertically into the flat top. They
seem to have a preference for the top, since about ninety
per cent of the nests were there. These bees, like their
predecessors, also carried water to moisten the hard clay
*One scoring finds the turrets sealed at the orifice, but in
all the specimens of ‘bis kind which I examined, the burrows
been usurped poses used by the wasp, Tryporylon clavatum.
The Ecology of a Sheltered Clay Bank 177
to make it workable; then they carried out mouthful after
mouthful of soft mud in the form of pellets. These were
used in the construction of the chimney until that was
of adequate size, after which the surplus material was
kicked aside.
By August 20 the males had disbanded and were no
longer seen about the premises. They had not gone far,
however, but were around on the other side of the house,
Testing, one each in the wild morning-glory blossoms
which they had found there. They quietly occupied the
floral cups for hours at a time, serving no good purpose
that I could see, but only usurping the rightful place of
the pollen-hunting mother, and often even fighting des-
perately to keep her ont—a perfect dog-in-the-manger.
At the end of August, despite several days of rain, fe-
males were still at work bringing in loads of pollen. By
September 3, several cold nights had caused them to lay
aside their virtue of early rising. They were abundant
and actively engaged at this late date. By the 6th or
7th, however, only about a dozen bees were still alive, or
at least in evidence about their business, some bringing
in pollen and others actually excavating. On September
12, with the preceding night very cold, no bees were to
be seen at work in the morning; at noon, one crept out,
and by three o’clock, two more ventured. But even at
this late date, eight males were seen as before, huddled in
the cup of the morning-glory flowers, where they had
Spent the night. These flowers had not closed for the
night,* and even at ten a. m., these males were in a tor-
aan
ight,
*It is unusual for morning-glory flowers not to close at n
but this actually happened late in the season when the —— were
Cold and dewey. These flowers which remained open gave
lent shelter to the bees.
178 Trans. Acad. Sci. of St. Louis
pid, dejected condition, heavily covered with dew. On
September 23 it was quite apparent that these insects
were being rapidly reduced in numbers by death. Several
dead ones were picked up from where they had fallen
beneath the mouth of their burrows, and there was no
way of ascertaining how many more dead remained
within. It is natural to think that females of such
species as this continue to work to the last, but on the
occasion of my visit on October 3, there were three
mother bees that were no longer exerting themselves on
home cares, or in any way trying to work, but appeared
to be simply idly waiting to be overtaken by death. In-
deed they lingered on beyond their just day, for even
their food-plants, the pollen bearing flowers, had almost
all ceased to bloom.”
While I was unsuccessful in determining the number
of nests made by each mother, the total number of nests
for the whole colony from year to year is of interest. In
1917, there were 62 nests; in 1918, 55; in 1919 the number
went down to 37, and in 1920, 49; in 1921, 51 and 1922, 40.
When one sees the enormous population of Anthophora
in 1921, many of which were trapped and perished, and
compares it with the 1921 Entechnia population, one
sees a vast difference in their rate of increase; in the
five years residence, one species has increased phenome-
nally, while the other has barely held its own.
The dates of emergence for the various years will be
treated under climate, and other details are elsewhere
given in these pages.
*At even a later date than this, however, on October 17, 1919,
one was seen to enter its ‘ene at Hematite, Mo., several miles
south of the clay bank.
The Ecology of a Sheltered Clay Bank 179
The Mining-bee, Anthophora (Antheméessa) raui.*
(S. A. Rohwer].
In 1917 I noticed among the turrets of Anthophora ab-
rupta, three which turned upwards, as indicated by ar-
rows in Fig. 9. The next year, there were five of these
in the mass, and, since this showed a very distinctive
digression, I concluded that this might be an advanta-
geous variation, and began forthwith to philosophize
upon the laws of variation, heredity, and survival of
the fittest, with the deepest solemnity. The reason why
the reader is mercifully spared this is that, when mak-
ing a later visit to the place, June 13 to 19, 1920, I was
Shocked to find about 90 of these upturned turrets, seg-
regated into three distinct colonies, and bees, which
greatly resembled A. abrupta, but still showed distinct
differences, industriously taking care of them! In all
characteristics excepting the upward turn, the turrets
were precisely like those of A. abrupta. Since these
bees had habits of digging, carrying water, and pro-
visioning their nests with pollen, all of which were so
very similar to the way of A. abrupta, it was not surpris-
ing that they proved to be taxonomically very nearly re-
lated to the latter species.**
Thus the great increase in the population in 1920
came practically as a saltation, for in 1918 they were
Practically nil. This sudden increase seems entirely pos-
sible, however, even though this species has the same
Short adult life eycle that A. abrupta has for I am of
the opinion that this bee makes more than one nest in a
“Described in Proc. Ent. Soc. Wash. 25: Sa 1923, by Mr. Rohwer,
who Rebbe that it is closely allied to sodal
brupta,
**In 1922, this species a peared three sas later than A. @
ated a few spaulava sues, seen a week after A. abrupta had all
off.
180 Trans, Acad. Sci. of St. Louis
season, and has three or four cells to each nest. By the
third week in July, the population was noticeably wan-
ing, and before the end of the month had almost dis-
appeared.
When these bees make their turrets, they use a process
which I have not yet been able fully to analyze. The
bee walks out backward, and when the tip of the abdo-
men touches the rim of the chimney, she applies her
pellet of mud. At first this is quite crumbly, but soon
one sees that in some way water passes over it, making
it more plastic and adhesive. One can see the water as
it spreads over the mud, but cannot see where it comes
from. It does not seem to come from out of the ab-
domen and if it comes from the mouth one cannot see
how it is passed to the tip of the abdomen. One load of
mud brought from within makes from one-fourth to one-
third of a ring.
‘As mentioned before, the only perceptible differences
between the chimneys of these bees and those of A.
abrupta, are that these turn upward instead of project-
ing from the bank horizontally, and these are usually
solid tubes while those of A. abrupta have a gap or fis-
sure, of unknown significance, extending the full length
of the tube on its upper side.* Sometimes this upward
turning chimney goes into pretty curves, but it never
fails to turn upward. This distinction is unusually short
and clear for the two species; I have never yet seen an
A. abrupta go into one of these vertical turrets, nor
have I ever seen an A. raui enter the horizontal ones,
although I have often seen both species go into holes
that had no turrets.
*Out of several hundred turrets of A. ebrupta, only twelve lacked
this split.
The Ecology of a Sheltered Clay Bank 181
My earlier notes show that in 1917 there were only
three of these upturning turrets, which indicated that
this species was newly arrived then. As early as June
18, 1921, I counted 40 such turrets, with A. rawi coming
in and out. They were not scattered over the whole bank,
but were distributed in groups; this probably means
that the bees build very near to the spot where they are
born. Fig. 10 shows a colony of 20 upturned turrets
in a small area (for this picture a wad of cotton was -
placed in each vertical chimney, to better distinguish
them from their neighbors), and Fig. 6 shows a mother
twice natural size.
I accidently broke off the turret of one nest about an
inch long, while it was in course of construction. The
bee coming out with the next load of mud did not seem
in the least surprised, but without turning about to ex-
amine the condition, she placed her load at the base and
began on a new turret. On June 18th ten new turrets
Were just begun.
The following record of one typical bee’s work gives
an idea of the manner of making the turrets. This bee
entered a shallow old hole and kicked out a small quan-
tity of loose dust; after some minutes of this, she left
and returned presumably with water. She then began
to bring out loads of mud and started to make her turret.
The face of the bank here was uneven and her first tine
loads went to fill the depression, first above the orifice
and then below it, until it was level and ready to re-
ceive the chimney, which she continued in the same man-
ner. A trip for water usually consumed two minutes.
In leaving the nest for water she came out head first,
having turned around in the tunnel; in coming out with
mud, she always emerged backwards and applied her
182 Trans. Acad. Sci. of St. Louis
mortar. Between 11:10 and 2:30, the burrow was dug
and the chimney built up to a height of % inch; from
that time until five o’clock, nothing more was done.
Like A. abrupia, these bees build very definite cups of
mud within the burrows, in which to oviposit. Fig. 11
shows a group of such cells—some of them opened to
show the pupae within, while Fig. 12 shows the pupae ,
in two stages of development. The interior wall of this
mud cup is varnished with some substance which renders
it waterproof, a condition we have already described for
A, abrupta.
Halictus bee, Halictus (Chloralictus) zephyrus Sm.
[J. C. Crawford].
A small burrow which went down vertically on the
top of the bank first attracted my attention in 1917. It
contained a pair of these Halictus bees. One of them,
probably the male, closed the aperture of the burrow
with his head, and prevented his consort from entering
until it suited his whim to admit her. This was the only
nest of this species found that year, and it now seems
likely that in taking the pair for identification, I exter-
minated this species in this locality, for in the two years
following none appeared.
The species must have been reinstated, however, be-
cause in 1920, three burrows were again discovered, and
in 1921, a dozen nests were there. They dug their bur-
rows horizontally into the face of the clay bank near the
base. Below each burrow was a little mound of pulver-
ized earth. It was at length discovered that several
adults occupied one gallery, and the nesting cells
branched off from either side of this gallery (Fig. 13).
The Ecology of a Sheltered Clay Bank 183
Rach of these cells, or pockets, contained an egg attached
to a small pellet of yellow pollen.
At first I thought that it was always the head of the
male which plugged the doorway; but as I later learned
that each burrow was really an apartment-house occupied
by many adults, I have been in doubt whether the one at
the door may not have been one of the feminine oceu-
pants. This common passage way is larger underground
than is the doorway, which is just wide enough to admit
one bee at a time. One nest which I opened had a
tunnel in the hard yellow clay nine inches long and three-
sixteenths of an inch wide, excepting at the narrow en-
trance. Pockets on either side of this tunnel had young
in all stages of development, from minute creatures cling-
ing to a little ball of pollen, to pupae already deeply pig-
mented. Besides these there were a half-dozen adults,
possibly mothers or adult sisters. After the nest had
been dug up, eight bees returning home congregated
before the ruins. The adults as they emerge nidify
in the same tunnel and gradually extend it. While these
are solitary bees they are very neighborly, and this
habit of community dwellings seems to point in the di-
rection of socialization. Indeed it almost seems to be
a link between the solitary and the social habit of bees.
Near the end of the season, October 2, 1920, and
again September 28, 1922, at Wickes, Mo., I witnessed a
new phase in the life of this species. Thousands of
these little bees were executing a sun-dance on the sunny
hilltop, where the grass was closely mowed. They were
in a great many groups of a few bees to several hun-
dred, while a few groups had several thousand. Some
groups were close together and some isolated, some dane-
ing over the short grass and others over the barren olay
184 Trans. Acad. Sct. of St. Louis
spots. Thousands were taken in the net, but most of
them escaped through the large mesh. They kept near
to the ground and moved rapidly; they were so small
that individuals could not be followed. Some burrows in
the grass were plugged with the heads of the guards
and in some the females were seen to enter. From: the
size of those in the dance, I suspected them to be males,
and the behavior indicated courtship.
Little Carpenter-bee, Ceratina calcarata Rob. [J. C.
Crawford].
A good many stems of the red rambler roses had been
eut back, and each year dozens of these contained nests
of these bees and their parasites, but in so far as I could
see there was no relation between these bees and the
other inhabitants of the clay bank, They went afar for
their pollen, and neither preyed upon the others, nor
were preyed upon; in fact, in every way they held them-
selves aloof from their neighbors.
The caterpillar wasp, Ancistrocerus fulvipes Saussure.
Among the pioneers in the clay bank should be con-
sidered the caterpillar huntress, Ancistrocerus fulvipes,
since she almost never used the old burrows for nest-
ing, but dug her own tunnels among those of the min-
ing bees.
In 1917, this species was seen about the clay bank as
early as June 17, when three adults were out. By July
7, this number had increased to a dozen, and by the
16th, neither their number nor their industry had waned,
although by this time, their companions Anthophora
abrupta, Trypozylon clavatum and the chalcid parasite
Monodontomerus had disappeared, either temporarily oF
permanently for the year according to species. For sev-
The Ecology of a Sheltered Clay Bank 185
eral days these mothers had been digging, carrying mud
or bringing in caterpillars. On July 30 I returned after
a few days absence and found the numbers of this wasp
materially reduced, so I suspected that this was about
the time for the passing away of the first generation.
Thereafter, a careful watch up to August 10 revealed no
trace of this citizen of the colony. On August 14, one
wasp, probably the first of the second generation, was
found at work. Up to ‘August 20, this one and another
were the sole representatives of their tribe, and on Au-
gust 30, three were out. Although September 3 and 4
were almost sunless, yet 7 individuals were active; this
was the maximum number of the second generation
Seen at any one time; from September 6 to 23, only
two or three at a time were out at work. On the last
visit of the year, on October 3, the day was gray and
chill, but it found two of these mothers still at work,
one plugging her burrow with mud and the other bring-
ing in caterpillars, both plodding faithfully on toward
the completion of their work, oblivious to the fact that
it Was now only a matter of days or perhaps hours until
for them the sun would shine no more.
In 1918 this wasp, like the other species described
above, emerged and began its work much earlier than
in 1917; on May 28, they were already out and busy.
In 1919, no records were made of their emergence, but
in 1920, when the bank was visited on May 27, they were
again found at work. Their appearance is correlated
- With temperature conditions in later pages. They ap-
peared in slightly greater numbers in 1918 than in 1917.
On July 17, about the same number were at work; on
July 31, none at all; hence this is evidently approxi-
mately the dividing line between the two generations. A
186 Trans, Acad. Sct. of St. Louis
visit just a month later showed a partial return of the
population.
No notes were made for 1919. In 1920, as already
stated, May 27, found these wasps already alive. Their
activity continued throughout June and the most of
July. I was unable to visit the grounds then, at the
date when the depletion of the first generation was ex-
pected, but on September 2, A. fulvipes were more
abundant than ever; 13 individuals were seen at one -
time, the greatest number yet seen at work simultane-
ously, which indicated that they were on the increase.
None were in evidence on October 3.
These notes, while not based on large numbers, are
probably sufficient to show an increase in numbers, an
earlier date of emerging when the weather is warmer
as with the other inhabitants of the bank, and a strong
indication of two generations a year.
These wasps, like the mining bees, went elsewhere
than the bank for the food for their young and them-
selves; hence the relation that this species had to the
community as a whole was in bequeathing their discarded
tunnels to the other inhabitants who might want them,
and in occasionally falling prey to other insects. One
such instance was observed on June 28, 1920, when a
bug, Reduvius personatus L. (W. L. McAtee), which was
amply protected in coloration by having its sticky body
covered with dust, and which was half concealed in a
crevice, was found feeding upon this wasp.
This eumenid was elsewhere seen to nest in tunnels
in logs; for an account of its life history, see ‘‘ Wasp
Studies Afield.’’
The Ecology of a Sheltered Clay Bank 187
The caterpillar wasp, Ancistrocerus unifasciatus Sauss.
[S. A. Rohwer].
Three mothers of this species were seen carrying
green caterpillars into cracks of the old building above
the clay bank, on May 28, 1921, where they were evi-
dently nidifying in the old mud nests of Sceliphron cae-
mentarium. I have previously recorded* the fact that
they use the old cells of this mud-dauber.
The spider-wasp, Episyron biguttatus Fab. [S. A.
Rohwer].
This Pompilid wasp was seen kicking dirt into a small
burrow at the base of the clay bank. She would quickly
Sweep in the earth with her front legs and then with
rapid beats of the tip of the abdomen she would pound
it down. The wasp was taken and the burrow opened;
one-half inch below the surface we found a perfectly
round cell of the size of a pea, which contained a spider
with an egg cemented to the dorsal side of the abdomen.
The pipe-organ wasp, Trypoxylon politum Say.**
This wasp should probably be included among the
. inhabitants, since three large pipe-organ nests were
Were built on boards or joists of the porch over the
bank, but a more logical and interesting reason for
including it was that several of these mothers were seen
foraging for spiders which made their homes in the old
bee burrows. Thus the species which seemed almost an
outsider could affect the life of the community even
though in a small way, by reducing the number of spider
*Ent. News 24: 396, 1913.
**Formerly called T. albitarse.
188 Trans. Acad. Sci. of St. Louis
occupants which in turn would have reduced the number
of bees and parasites. One nest gave forth its adults
between June 26 and 29, 1917.
The mud wasp, Sceliphron caementarium Drury.
This mud-dauber stands in the same relation to the
life of the unit as does the pipe-organ wasp, just men-
tioned. These yellow-legged wasps, too, plastered their
mud huts on the wooden porch overhead, and in one in-
stance even attached one direct to the clay bank, in a
depression which I had made in digging out a nest; a
crooked, lop-sided, distorted piece of work it was, and
it could not have been different in the cramped corner,
but with yards of flat surface overhead, why should a
sensible wasp choose a spot of this kind!
Like the Trypoxylon politum, these wasps were occa-
sionally seen entering the burrows in quest of spiders.
Their discarded nests about this unit were utilized as
homes by Trypozxylon clavatum, Ancistrocerus wnifas-
ciatus, and Pseudagenia mellipes and the burglar wasp
Chalybion caeruleum made the best of the opportunity
to break into them in a most desperate way.
The Larrid wasp, Tachysphex terminatus Smith.
These little wasps appeared about July 16, 1917. They
were digging their burrows in the loose dirt on the top
of the clay bank and provisioning their nests with short-
horned grasshopper nymphs. Toward the end of the
month their numbers became much reduced; on July 30,
only three females were in evidence.
These wasps are supposed to be essentially sand-
loving, but here we found them digging in the loose dirt
The Ecology of a Sheltered Clay Bank 189
on top of the hard clay bank, and as a consequence the
nests were very shallow, so shallow, in fact, that the prey
could be exposed to view by merely blowing the dust
away, whereas those burrows found in sandy areas had
a much more substantial depth.
On August 10, I found that the number nesting had
increased to six, which would suggest, though not prove,
a second generation. Also it became evident at that
time that this wasp sometimes falls prey to the other
inhabitants; one dead specimen was taken from the web
of a spider occupant of the bank. Only four days later
I found these Larrids had suddenly increased to about
thirty, with a goodly proportion of males in the lot.
Excited courtship and mating were the program of the
day. The males outnumbered the females, and the usual
fight for partners occurred, three or four males some-
times struggling together for the possession of a fe-
male. Even during this commotion, however, about ten
females managed to attend to home duties, between inter-
ruptions, bringing in locust nymphs mostly of the spe-
cies Chortophaga viridifasciata De G. (A. N. Caudell)
(Fig. 14). Toward the end of August the number of
mothers began to diminish; on September 3, one was
found dead in its doorway;-this probably indicated the
approach of the season of natural death. And so it was,
for only one or two were seen after that, and on my
farewell visit for the year, Oct. 3, none of these sun-
loving creatures were found to have survived the cold,
gloomy days just preceding. The wasps have never
appeared in early spring, and a few cocoons taken from
the bank on April 28, 1921, gave forth adults on June
2; this indicates that this is a summer and fall creature.
In 1918 these wasps appeared in slightly greater num-
190 Trans. Acad. Sci. of St. Louis
bers than in 1917; by 1920, the population had trebled.
Just why this habitually sand-digging Larrid should
choose to adopt itself to the yellow clay of the bank, and
why after this change, it should flourish and increase
in numbers, is still an unanswered question.
We see only slight relation between this wasp and the
other inhabitants of the clay bank, since these mothers
must go abroad for the Orthopterous prey; hence prob-
ably it had nothing in common with its neighbors other
than occasionally falling prey to the spiders there.
The devil’s horse, Stagmomantis carolina Linn.
The egg-cases of the devil’s horse were often found
plastered to the boards above the bank, and each June
the little nymphs could be seen walking about the bank.
They are carnivorous, and probably fed upon some of
the very small insects about the bank, until they in turn
fell prey to spiders that were occupying the old burrows.
The paper wasp, Polistes pallipes Lepeletier.
During the four years there were probably a dozen
nests of P. pallipes on the under side of the porch and
attached to the rose bushes in front of the bank; but in
so far as I could see, the wasps had no relation to the
unit as a whole, since they. made their own nests, and
did not get prey from the clay-bank; neither did they fall
prey to the inhabitants. Sometimes, they built their nest
in narrow spaces between boards, which resulted in nests
strangely shaped, but the point which remained most
puzzling was that every year some should persist in
building in this cramped space between two joists, and
making these unusual nests, when there were hundreds
of square feet of clear ceiling and wall space ready to
accommodate them.
The Ecology of a Sheltered Clay Bank 191
The stick-bug, Emesa brevipennis Say [W. L. Me-
Atee].
Another insect which should be classified as a pio-
neer in the unit, even though it makes no nest, is the
stick-bug, Emesa brevipennis. It spent its whole life
among the rubbish, boards, ete. piled on top of the
bank. The nymphs were seen as early as July 16, and
the adults as late as October 12. During their im-
mature stages they were inconspicuous, hence not fre-
quently noticed, but when full grown and more easily
noticeable, they were observed in great numbers. Since
they are carnivorous in habit, they were probably very
directly responsible for some changes in the popula-
tion of the unit. While I have never chanced to see
them actually prey upon the inhabitants of the bank,
I have seen them feeding upon insects which had
been visiting the bank for shelter. One was imbibing
the juices of an adult moth, Pyralis cuprealis Hubn. (H.
G. Dyar); another was seen to hold an immature (sec-
ond stage) short-horned locust in the fore-legs, mantis-
fashion, and, with its beak inserted, imbibe its juices
for a whole day. Whether they actually captured sleepy
bees and wasps, I cannot say.
‘Ants. Crematogaster lineolata Say [W. M. Wheeler].
A crack in the porch post was the home of a large
colony of these ants. Its part in the game of give and
take being played on this scene was to supply food to
the ant-lion larvae, whose pits were in the dust at the
base of the bank. The spider, Habrocestum pulex, and
also an unidentified ant-mimicking spider were observed
feeding upon them. These ants often covered up the
cracks in their posts with a gummy secretion. On Au-
192 Trans. Acad. Sci. of St. Louis
gust 14, 1917, many winged specimens were in the lot.
This colony remained a permanent part of the poput-
lation during the five years observation.
On my spring trip afield, April 2, 1921, I found on the
ground about 400 of these ants, dead. They were still
soft; hence they could not have been dead long. More-
over, they had not been there on February 26, when
I examined the bank. I had previously noted these ants
hiding in the cracks in these walls and plugging the
crevices with a gum-like substance; hence they could
not have merely fallen out. The question remains
whether these had come out to meet the spring and had
been caught by a severe frost, or whether they were the
accumulated dead of the winter, which had been dumped
out by the survivors.
In another instance, when a Polistes pallipes queen was
taken from the nest above the bank for use in homing
experiments, a colony of these ants discovered the nest
soon after it was left unguarded; they bit into the walls
of the cells and completely removed the larvae bit by bit.
This task consumed two days, during which time the
nest was black with ants.
Anti-lion larvae, Myrmeleon immaculatus De Geer
{A. N. Candell]. Myrmeleon mobilis Hagen [A. N. Cat-
dell].
The numerous insects mining in the bank kicked out
quantities of finely pulverized clay, which accumulated
at the foot of the bank (Fig. 1,X). This made an excel-
lent abode for the ant-lion larvae, which dug their pits
of various dimensions (Fig. 15) and fell heir to occa-
sional bits of provender brought in for the other nests
but dropped, as well as sometimes the spider or ant
The Ecology of a Sheltered Clay Bank 193
inhabitants themselves or occasional visitors to the set-
tlement. That this was a favorable place for them to
flourish and multiply was evidenced by the fact that in
1917, 20 pits were counted at the foot of the bank; in
1918, 150 were there, but most of these were gathered
to send to a friend for study. Hence in the following
years they appeared again in about the same numbers
as in 1917, at the base and also on top of the bank. Of
course we should bear in mind that the ant-lions, by
virtue of their peculiar mode of development, are well
prepared to survive and flourish, so that, like the poor,
we shall have them always with us. They neither perish
nor migrate if food becomes scarce for a time, but they
simply lie dormant, and put off growing and transform-
ing for a year or so until food again becomes plentiful.
Thus, an examination on October 13 revealed many
ant-lions in all stages of development; some were re-
constructing their pits in my presence, others were
larvae in all sizes from small to large, with pits in pro-
Portionately varying sizes. Precisely these conditions
were found at all other seasons, from the opening of
spring to the close of autumn. This shows that they
have no set time of the year for their metamorphosis,
but change whenever they have reached a certain stage,
regardless of the time of the year. Some that were
brought into the house and fed abundantly on flies
8rew rapidly, pupated in a few days and emerged as
adults from three to four weeks later.
One must not forget to consider the life habits in rela-
tion to the environment, when contrasting the develop-
ment of the ant-lions with that of the bees. The larvae
of the bees are supplied with sufficient food to carry them
through their development without their exertion, while
194 Trans. Acad. Sci. of St. Louis
the ant-lion larvae must get what they can. With the
ant-lions, it is either a feast or a famine—when food
is plentiful, they develop to maturity quickly; when it
is scarce, they do not die, but combat famine with pa-
tience. Herein Mother Nature anticipated Victor Hugo,
who said: ‘‘A clock does not stop short at the precise
moment when the key is lost!’
An interesting detail in their method of feeding was
noticed when a Grapta caterpillar was dropped into one
of the pits. The ant-lion captured it and promptly
sucked out the juices; five minutes later it was discov-
ered that the ant-lion had also snipped the body-wall
and was dragging the viscera out through the aperture.
The only relation these Jarvae had to the other inhab-
itants of the unit, was to eat any insects, such as weevils
and caterpillars, which, through accident or misfortune,
fell into their pits. I noticed also that occasionally their
pits served as good starting points for the burrows of
the Larrid wasp, Tachysphex terminatus. The Antho-
phora bees were probably too big game for them, but 1
did find one struggling with a dead bee in the pit.
Whether the dead bee had fallen into the pit, or the ant-
lion had actually attacked and killed a live one which
eame within its reach, I do not know.
One ant-lion passed five weeks in the pupal stage; hav-
ing spun the cocoon on ‘April 16, 1921, it emerged as
adult on May 24, A second one spent six weeks in that
stage, or from April 6 to May 25, 1921.
House-Spider, Theridion tepidariorum Koch [J. H.
Emerton].
These house-spiders should be reckoned as among the
original inhabitants of the unit, since their webs were
The Ecology of a Sheltered Clay Bank 195
occasionally found about the porch above the bank. Their
actual activity was observed only once, on October 12,
when one was intermittently feeding upon a large, hairy
caterpillar caught in its web. I feel confident, however,
that, since they are night prowlers, they participated
more fully in the life of the community. Elsewhere I
have found this species feeding upon a wasp (Odynerus
anormis Say).
Résumé of Pioneers.
To summarize, then, the five years’ work at the clay
bank gave us ag pioneers, those which actually opened
the way for others, the eighteen species listed below. It
is interesting to note that six of this number were soli-
tary bees, seven were wasps, and the remaining five be-
longed to as many different orders.
Carpenter bee. Xylocopa virginica.
Mining bee. Anthophora abrupta.
Mining bee. Entechnia taurea.
Mining bee. Anthophora raut.
Halictus bee. Halictus zephyrus.
Small carpenter bee. Ceratina calcarata.
Mining wasp. Ancistrocerus fulvipes.
Eumenid wasp. Ancistrocerus unifasciatus.
Spider wasp. Episyron biguttatus.
Pipe-organ dauber. Trypoxylon politum.
Mud-dauber. Sceliphron caementarium.
Larrid-wasp. Tachysphex terminatus.
Paper wasp. Polistes pallipes.
Devils horse, Stagmomantis carolina.
Stick-bug. Emesa brevipennis.
Ant. Crematogaster lineolata.
196 Trans. Acad. Sct. of St. Louis
Ant-lion larvae. Myrmeleon mobilis. & M.
ammaculatus.
House spider. Theridion tepidariorum.
(c) Renters.
In this chapter are enumerated the secondary inhab-
itants of the old clay bank, those which came as ten-
ants of the burrows made by some of the creatures
listed in the foregoing chapter. These, for the most
part, were less hardy than the pioneers; they came into
the community, evidently quite willing to utilize its ad-
vantages and share its comforts, Often they dropped
in merely for shelter or food, and remained to the end
of their days. These renters, besides using the old dwell-
ings, influenced the population of the unit in two ways
—by using the other inhabitants for food, and often by
being eaten by them.
The Pompilid wasp, Pseudagenia mellipes Say.
This wasp was occasionally seen about the clay bank.
There were four in 1917 and perhaps a dozen in 1920.
In ‘‘ Wasp Studies Afield’’ we have told how this crea-
ture nests in both new and secondhand domiciles; some-
times she makes small mud nests under loose bark (one
such nest was found inside an oak-apple lying on the
ground), and at other times she modifies the nests of
the mud-daubers to suit her needs.
The interest of P. mellipes in the bank was the prey
which it sought there. The wasps were often seen going
into the old bee-burrows, now occupied by spiders, prob-
ably in search of food for their young. On a few occa-
sions, they were seen going elsewhere for prey and
The Ecology of a Sheltered Clay Bank 197
bringing home their booty. The legless spider brought
in by one of these P. mellipes was identified as Pisaurina
undata Htz. [C. L. Shoemaker] and on another occa-
sion a mother was seen at the base of the bank carrying
a half-grown Phidippus spider. This versatile little wasp
found in the old bee-holes both home and hunting-
ground; she was actually seen to drag a spider up out
of one burrow—its lair—and down again into another
hole near by—her own nest. Thus we see this citizen
finding in the clay bank ‘‘all the comforts of home,’’
shelter or a domicile already provided, and food sup-
plies available near her door, and she has just as little
work to acquire the one as to get the other, whereas in
the usual condition, the home-makers get the benefits of
one or the other, but seldom of both. These wasps con-
tinue their activity throughout a fairly long season, from
the last of May to the first of September.
Since they made their own mud cells in the old bee
burrows, they are classified with the renters, although
they had just as much claim to be among the pioneers.
Trypoxylon plesium Roh.* [S. ‘A. Rohwer].
Only one specimen was seen here, and it was taken at
the clay bank on July 7, 1917. I saw this wasp actually
walk over two distinct spider webs without becoming in
the slightest way entangled; evidently it was on a spider-
hunt when captured.
es
198 ' Trans. Acad. Sct. of St. Louis
Trypoxylon clavatum Say. [S. A. Rohwer].
This species of wasp in which the male parent guards
_ the nest while the female hunts, has been recorded as
making use of old beetle burrows in logs, and cleaning
out and using the old cells of the mud-dauber wasps. In
the clay blank, they were quite abundant, here using the
empty bee burrows, partitioning them with mud and
filling each compartment with spiders.
When this species first made its appearance in 1917,
about June 28, there were only three nests, each with its
male guarding the doorway, and four days later there
were seven. By the time of the next visit, on July 16,
these had vanished, and for two weeks not a single
T. clavatum was to be found. On July 31, the first one
of the second generation appeared. Time dragged on and
this one continued to be the sole representative of the spe-
cies. Not until the first of September did others appear,
and during that week a half-dozen or more were at work.
During the latter half of that month, they succumbed to
the inevitable. While 7. clavatwm appeared in the fol-
lowing years, their numbers became fewer; the species
was not holding its own. No doubt one very consider-
able factor in the reduction of their numbers was the
destruction I wrought in digging up their nests when
their colony was not yet strong, in order to study their
prey.
The relation of these insects to the unit was pri-
marily in using the old burrows; with the male on guard
in each nest it is not likely that the young of this wasp
serves as host to any parasites. A comparison of the
contents of their nests with the list of spider inhabitants
of the clay bank shows that they must have gone afield
The Ecology of a Sheltered Clay Bank 199
for their prey. These details of the life history will be
presented in a later publication.
Trypoxylon albopilosum Fox [S. A. Rohwer].
This wasp, with habits similar to those of T. clavatum,
likewise oceupied the old burrows of the mining bees.
It was first observed on June 28, 1917, when several
mothers were at work. They always appeared only in
such small numbers, however, that I could not accurately
define any seasons or generations for them, only that
they were to be seen occasionally until early Septem-
ber. Since I opened part of the nests to study the con-
tents, it was only natural that their population dwindled
away, until in 1920 one lone survivor was observed, The
relation of this species to their neighbors was the same
as that of T. clavatum.
The Monobia mud-wasp, Monobia quadridens. Lin-
naeus.
This wasp, commonly called the carpenter mud-wasp,
but, according to our account in ‘‘Wasp Studies Afield”’
not a carpenter at all, can be regarded as only a sec-
ondary inhabitant of the clay bank, since it made no
nest of its own, but occupied that of the carpenter-bee,
Xylocopa virginica. These wasps were seen each year
bringing in mud for partitions and caterpillars for food
for their young. On one occasion, as previously re-
corded, one Monobia mother actually utilized the tunnel
ofa mining-bee, carrying in caterpillars and sealing up
the opening with mud.
This wasp appeared from year to year in very limited
numbers, not more than 6 or 8 being present at work at
200 Trans. Acad. Sci. of St. Louis
any one time, and most of the time not even that many.
Since the shedding-skins of the silver-winged parasite,
A. tigrina, were often to be seen adhering to the wood
near the holes, it seemed probable that Monobia suf-
fered its share of the ravages of this pest.
Their season was long; the first appeared about June
26th, 1917, and on October 3, two were seen, still cling-
ing to the woodwork over the bank, stunned and half
dead with the cold, while a third one was seen to enter
its tunnel. In 1918, they appeared in decreased num-
bers, and by 1920, only three could be found. I sus-
pected that A, tigrina was more responsible than any
other factor for their elimination.
The grass-carrying wasp, Chlorion (Isodontia) au-
ripes Fernald. [S. A. Rohwer].
This wasp, a very conspicuous if not numerous mem-
ber of the unit, occupied the old tunnels of the carpenter
bee, and occasionally she used the old burrows which had
been made in the clay bank by the mining bees. It 1s
very easy to tell just where this wasp mother has her
nest, for instead of plugging the opening with mud or
otherwise concealing it, she stuffs the aperture with
grass, drawing each strand in by its middle, so that a
broom-like tuft protrudes prominently, thus revealing
at once to an experienced eye the location of her nest.
This insect occurred in very modest numbers; no more
than three or four individuals were at work at any one
time. In 1920, neither the wasps nor their nests were
to be found there. There was evidence to lead us to
think that the silver-winged parasite had played havoe
with these wasps, the same as with the other occupants ©
of the carpenter bee galleries.
The Ecology of a Sheltered Clay Bank 201
While these wasps, as well as the Monobia mud wasp,
were to be regarded as secondary occupants of the car-
penter-bees’ burrows, the relationship was more com-
plex, since by occupying these burrows they took just
that much space from the carpenter-bee, because each
generation of Xylocopa used the old domiciles, in so
far as they were adequate, and if the tunnels had been
usurped by other species, the mother carpenter-bees
were obliged to spend their time and energy at the slow
process of hewing wood instead of being fruitful and re-
plenishing the earth. Therefore, the only direct bearing
that these two wasp occupants had on the inter-relations
of life in the unit as a whole, was to deprive the car-
penter-bee of some of her rightful nesting-places and
to serve as a host for the parasite, Argyromoeba tigrina,
for they both gathered their food elsewhere.
The blue mud wasp, Chalybion caeruleum Linnaeus.
This wasp was heretofore regarded as the mud-
dauber wasp making nests very similar to those of the
“tyellow-legs.’? On the contrary, I have shown else-
where that this wasp does not make nests of her own,
but occupies the nests of Sceliphron caementarium,
either by breaking into a ‘‘live’’ cell and destroying the
Prey or by using the abandoned cells. In so far as the
bank was concerned, however, this species was occa-
Sionally seen foraging for spiders among the burrows.
On several occasions when one was foraging among the
Spider webs, she broke through or became entangled,
whereupon, in a very skillful manner, she quickly dis-
entangled the web from her person. Elsewhere, how-
ever, I have often found dead specimens that had been
caught in spiders’ webs.
202 Trans. Acad. Sci. of St. Louis
These wasps were found to be present at the bank in
great numbers on May 27th and 28th. This was before
their nesting activities had begun. The significance of
their presence has not yet been determined, so I could
only state the circumstances. They were on an elm
sprout six feet tall, which grew between the rambler
rose and the front of the bank. Other insects were there
too, so I thought possibly the aphids, which were plen-
tiful there, constituted the attraction, but I could not see
that they got anything from the aphids; in fact, the
plant-lice were on the under side of the leaves while the
wasps always walked about on the tops of them.
The wasps were constantly active on the shrub, flying
and hopping about, running nervously over the foliage
and jumping or dropping from leaf to leaf. The per-
formance began at about 7:30 a. m. when the first few
arrived and became active; the number rapidly increased
as time passed. Standing a short distance away, one
could see others coming in from various directions to
join the dance—for apparently this gathering was some
sort of social function—a stag affair, however, for @
dozen or so taken in the net proved to be all males. I
suspect that they were awaiting the emergence of their
mates, and that if I had been on hand at the proper
time, a little later, I could have witnessed some pretty
courtship festivities.
Leaf-cutter bee, Megachile campanulae Rob. [J. ©.
Crawford].
One specimen of this bee was seen at the clay bank;
it was at once taken for identification, and therein the
prospective founder of a colony was evidently destroyed,
since in the following years this bee was not seen again.
' Lhe Ecology of a Sheltered Clay Bank 203
It was on July 30 that this individual entered one of the
burrows. When this hole was opened, a quantity of the
resinous material was discovered, but I had no way of
knowing whether this or another insect had done the
storing. The Megachile bees are leaf-cutters, and make
nests in hollow stems, but I have elsewhere* recorded
that they have been known to make their leafy cups in
sheltered places as under clods of earth. Some bees
gather resinous substances, but the Megachiles are not
known to do so.
Another Megachile bee, M. generosa was taken while
at rest on the bank June 3, 1921, but it is unlikely that
its presence there was anything but accidental.
The Osmia bee, Osmia lignaria Say. [S. A. Rohwer].
This little bee did not appear in the community until
1920. In earlier researches I have found that this spe-
cies builds its nests in mud nests of Sceliphron caemen-
tarium. Since these bees could not offer the excuse of
foraging as an explanation for their presence here, I at
once suspected that they were replastering the old bur-
rows of the other bees here for their own nests in the
same way that they utilize the old mud-daubers’ cells.
This I soon found to be true. ‘About a half-dozen of
them were seen thus occupied about the bank from May
28 to June 19, 1920, and a month earlier in 1921. They
Were also observed gathering pollen from the black-
berry blossoms near by.
Spider, Ariadna bicolor Htz. [C. L. Shoemaker].
Of spiders, this was the most abundant species among
the inhabitants of the clay bank, and from the first of
— ey
*Trans, Acad. St. Louis, 24:1-71, 1922.
204 Trans. Acad. Sci. of St. Louis
September until frost their webs became more and more
conspicuous. They made funnel-shaped webs, with the
points of the funnels extending deep into the old bee
burrows, and got a good living from the small fry that
became entangled in the webs, such as visiting snout-
beetles, ants, chalcids, flies, ete. I was surprised to find
this spider actually feeding upon the bug, Reduvius per-
sonatus Linn. [W. L. McAtee]. Occasionally, however,
the spiders themselves fell prey to some predatory
wasp.
Spider, Steatoda borealis {[J. H. Emerton].
Among the most important renters of the old burrows
in the bank were the spiders, Steatoda borealis. They
made light, irregular snares about the bank, more often
about the old burrows, where many of the smaller in-
sects became their prey. I have often seen Chalcid para-
sites and ants, Crematogaster lineolata, entrapped in
their snares. The species may rightly be regarded as
an inhabitant of the bank, either a renter or a pioneer.
They were very numerous in 1917, and only moder-
ately so in the following years. They were as abun-
dant as ever even so late as October 3, when one spider
was making several brave attempts to subdue a large lo-
cust, Tettigidae lateralis var. polymorpha Burm. (A.
N. Caudell), which was entrapped in its web. Two un-
identified beetles were also their victims that day.
Résumé of Renters.
The following table summarizes the renters in this
social group. They number eleven species, of which
seven are wasps, two are bees, and two are spiders. All
The Ecology of a Sheltered Clay Bank 205
of these utilized to good advantage the burrows or the
nests left by the pioneers, and they in turn left many of
their burrows for a third tenant. Among these renters
one might also include the bees Anthophora abrupta, be-
cause they sometimes used their old burrows for a sec-
ond season, after enlarging and renovating them.
Renters of Old Burrows.
Pompilid wasp,
Spider wasp,
Spider wasp,
Spider wasp,
Monobia mud-wasp,
Grass-carrier wasp,
Cow-bird wasp,
Pseudagenia mellipes.
Try poxylon plesium,
Try poxylon clavatum.
Try poxylon albopilosum.
Monobia quadridens.
Chlorion auripes.
Chalybion caeruleum.
Leaf-cutter bee, Megachile campanulae.
Osmia bee, Osmia lignaria.
Spider, Ariadna bicolor.
Spider, Steatoda borealis.
(d) Visitors.
This is a list of insects and larger animals which came
into the community quite by accident, or in quest of food
or shelter. In some cases they remained as permanent
additions to the group, but in most cases, their stay was
temporary. Many of them influenced the lives of the pio-
neers and the renters in one of two ways: either by using
them as food, or by giving themselves as food.
Whether the word visitor or transients would be the
better term to apply to this group is uncertain. Many
insects alighted on the bank quite accidentally; others
came there for the definite purpose of securing prey;
*%
206 Trans. Acad. Sci. of St. Louis
some which came as visitors remained at the bank to
nest. It is this latter group which prevents one from
definitely drawing the line between visitors and resi-
dents; the parents of all the occupants must at one time
have been visitors, but not all visitors were occupants.
Reptiles.
On October 3 I noted a snake (Fig. 16) at:the base of
the bank, with a small portion of its body still within
the rodent burrow, which I suspected was its home. It
lay all day long in one position, as figured, but escaped
when I attempted to capture it. At that season of the
year, with food scarce, it probably had to wait a long
time for a morsel, but if this creature had its home at
the base of the clay bank all summer, it was no doubt
a considerable factor in the control of the insect popula-
tion during the season.
Common lizards or swifts.
During the entire summer of 1917 several common
gray lizards made their home among the piled wood on
top of the bank, and were undoubtedly a factor in reduc-
ing the life of the group. They, too, had enemies, prob-
ably larger ones, which I was unable to discover, and
these all attacked the lizards in the same way, in the
cases observed. During the summer, three dying lizards
were found there at different times, all with torn ab-
domens and entrails protruding, as shown in Fig. 17.
They were always found in the early morning hours,
which indicated that the attack had been made during
the dark hours of the night.
It was strange that these gray lizards did not appear
The Ecology of a Sheltered Clay Banh 207
the following season, although they were abundant along
the railroad tracks less than two hundred yards away,
but instead the blue-tailed lizard or skink appeared in
their place. The blue-tailed lizard had evidently sup-
planted its predecessor, but since its habits are the same,
very probably its function among and relation to the
inhabitants and visitors at the bank were practically
the same. This idea was strengthened when on three
mornings in early June, 1922, a third species Cnemido-
phorus sexlineatus [D. M. Cockran], was seen coming
out of a burrow and calmly making the rounds of the
clay bank, often entering the bee burrows. It would
enter head first, with the tail protruding, and often it
would perform the quaint feat of, twisting its tail about
the turret while its body was inside the tunnel. It
seemed to find the burrows adequately roomy, for it
would enter head first and also emerge head first. What
it got out of the burrows I do not know, but since this
lizard was seen to snap at Anthophora bees in the open,
I suppose many of these bees were eaten within the
burrows. The Chalcid parasites were also in abundance,
and wild roaches were often seen in the burrows. The
red-eyed flies, Ganperdea apivora, were numerous
at just that time, and the skinks were seen successfully
capturing these, and a few undetermined insects, in the
open.
Toads.
In 1920 a big toad was a familiar figure about the
bank. Before the end of the season, he had the portly
figure of a war-time profiteer, and we felt justified in
Suspecting that the rest of the population had suffered
accordingly.
208 Trans. Acad, Sci. of St. Louis
Phoebe bird.
Among the vertebrates which were at home at the clay
bank was a phoebe bird and her annual family.’ The
nest, on one of the posts, was removed each winter, and
each spring a new nest appeared in the same place. One
year I noted two broods of young in the nest. Whenever
I was at work at the clay bank, the mother bird went
afar for her food, but often when I came suddenly upon
the scene, I saw her quickly fly away from among the old
lumber on top of the bank. From this I suspected that
she lost no opportunity to get her insect food near at
hand when she was not disturbed.
Red bird.
A red bird nested in the rambler bushes in front of the
bank, during the summer of 1920, but in so far as I
could see had no influence on the insect colony.
Ground mole.
There were a few rodent tunnels under the ground at
the foot of the bank, and one year there were two distinct
openings, but whether the moles or field mice affected
the insect life of the bank I cannot say.
Man.
The human inhabitants of the dwelling whose porch
covered the bank were not in the least a factor modifying
this unit; further than the first construction of the site,
no relationship existed. The house was almost always
vacant except at week-ends, and even then none of this
biped fauna came near enough to the bank to influence
the life, since they mistook the inhabitants for bumble-
The Ecology of a Sheltered Clay Bank 209
bees, and hence kept at a discreet and respectful distance.
However, one man was a factor to be reckoned with in
considering the enemies of the unit—that was the ob-
server. That he should rightly be considered as a non-
resident part of this ecological unit is evidenced by the
fact that, in taking specimens for identification, he dis-
rupted the fine balance of nature, but actually no more so
than if he had been a lizard and captured these insects for
food. He took only those species which were unknown to
him, and then took them in minimum numbers. However,
in some cases he was guilty of swinging the balance to
some point where, without his interference, it would not
have stood. Had he not taken the only Halictus, Mega-
chile and others, these insects probably would have be-
come established in the bank and their progeny would
have done their part to modify the life of the community.
The reader can easily estimate the extent of these depre-
dations by glancing over the list of the inhabitants and
noting all the species that have been submitted to ex-
perts for identification. I do not want to exaggerate
this point absurdly, but I think it is well that we should
realize that man is not a thing apart from the other fac-
tors in the balance of nature, but that he is only one
species among many in the great game of give and take
which makes up ecology.
Lepidoptera.
Pyralis cuprealis Hubn. (H. G. Dyar]. This visitor
was found in the beak of a stick-bug, Emesa brevipennis,
on July 16, 1917.
Two caterpillars belonging to the Pyralididae. [S. B.
Fracker] were found promenading on the bank on Sep-
tember 12, 1917, and later in the day a dead one was
210 Trans. Acad. Sci. of St. Louis
taken from an ant-lion’s pit. Thus, even though they
came to the place quite casually, they served the com-
munity.
Pseudaglossa lubricalis G. [H. G. Dyar]. One speci-
men was picked up dead at the bank on September 3,
1917
A caterpillar belonging to the Noctuidae [S. B.
Fracker] was feeding on a small plant growing at the
base of the bank.
Catocala innubens Guen. [Ernst Schwarz]. During
the latter part of July, 1918, several of these moths
sought daytime shelter here.
Herculia olinalis Guen. [H. G. Dyar]. The caterpil-
lar of this moth was found walking on the clay bank on
April 28. On the next day it spun its silken cocoon
and emerged as an adult on May 22.
Hymenoptera.
Pompiloides sp. [S. A. Rohwer]. This wasp was forag-
ing for spiders in the old bee tunnels during August and
early September, 1917.
Arachnophoctonus ferrugineus Say. One specimen
foraging for spiders on July 30, 1917.
Pompiloides americanus Beauv. [S. A. Rohwer].
This had come to the bank as a visitor on a foraging eX-
pedition, but had become a victim to the prey which she
was pursuing and her lifeless remains were removed
from a spider web on July 30, 1917.
Tachytes peptictus Say. [S. A. Rohwer]. On July
21, 1920, two or three of these green-eyed wasps were at
rest on the vegetation at the edge of the bank.
The Ecology of a Sheltered Clay Bank 211
Stlaon sp. (S. A. Rohwer) one was nervously walking
about the clay bank on July 30,
Sphea (Ammophila) nigricans Dahl. [S. A. Rohwer].
This was seen walking about the clay bank, August 14,
1917, evidently seeking a place for a nest. Since her
prey is Lepidopterous larvae, we cannot accuse her of
having come to the bank for prey.
Sphea (Ammophila) procera Klug. [S. A. Rohwer].
On August 16 this large wasp was seen burying a large
caterpillar of the Noctuidae [S. B. Fracker]. Since she
left her young ‘‘on the door step’’ of the bank, she should
be regarded as an inhabitant or a pioneer; but since I
took possession of both, she was thus reduced to a visitor.
Odynerus (Stenancistrocerus) unifasciatus Say. [S.
A. Rohwer]. This Eumenid was seen moistening a spot
on the bank with water and biting out and carrying away
the mud, June 28, 1920.
Polistes variatus was foraging on the rose bush in
front of the bank on May 28, 1920. There was hardly
any connection or rather only a far fetched one, in the
relations to the inhabitants of the bank, in that the P.
variatus would probably carry off caterpillars which, in
the natural course of events would have become food for
the inhabitants.
Hoplisus (Pseudoplisus) phaleratus Say. [S. A. Roh-
wer]. This species, seen entering a crevice on July 21,
1920, was distinctly new to the bank.
Trypoxylon nigrellum Roh. [S. ‘A. Rohwer]. One was
seen foraging for spiders in the old bee tunnels of the
clay bank on September 6, 1917.
Miscophus americanus [S. A. Rohwer]. Two females
taken from the bank on August 31, 1918, were probably
seeking a nesting site.
212 Trans. Acad. Sci. of St. Louis
Vespa germanica Fabricius. In August, 1920, one
worker of this hornet visited the bank at short intervals
for three whole days, and as described elsewhere, car-
ried off each time a specimen of the Chalcid parasite,
Monodontomerus.
Pseudagenia architecta Say. The first and only speci-
men of this wasp was observed on July 7, 1917, to
enter many old burrows of the mining-bees. Its be-
havior was very much like that of its cousin, P. mellipes
when hunting; hence we suppose that it was foraging.
This wasp makes tiny twin cells of mud that are some-
what barrel-shaped.
Megachile generosa Cress. [S. A. Rohwer]. This
bee always returned and rested on the bank after
{ missed capturing her on June 3, 1922, which indicates
that her interests were probably other than being merely
a visitor.
Halictus pectinatus Rob. [S. A. Rohwer]. ‘A male bee
of this species was found dead on the bank, September
6, 1917.
Photopsis sp. One individual was making itself at
home on the bank. This wasp is supposed to be para-
sitic; it had evidently found a host among the population.
Camponotus herculeanus L. subsp. pennsylvanicus
De G. var. ferrugineus Fab. [W. M. Wheeler]. These
ants, many of them workers and some with large heads,
were going about the bank, often entering burrows. They
first appeared in 1921.
Beetles.
Corymbites inflatus Say. [E. A. Schwarz]. This click-
beetle was at rest at the foot of the bank on July 16.
This beetle, or others of its kind, arriving there would
The Ecology of a Sheltered Clay Bank 213
probably have no effect upon the inter-relations of life
except as they might fall prey to some of the spiders or
vertebrates.
Photinus pyralis Linn. [E. A. Schwarz]. Many of
these ‘‘lightning-bugs’’ spent their days near the clay
bank about the first of August. Since they are nocturnal
insects and are in a sleepy condition during the day, they
should have been easy prey ‘to alert prospectors. No
actual observations were made, however, on the effect
of their presence upon the others.
Hypera (Phytonomus) punctata Fabr. [E. A.
Schwarz]. Two specimens were taken at two distinct
places on the clay bank on July 30, 1917. There may
have been more, but their coloration is such as to make
them almost indistinguishable from their surroundings.
They evidently fell prey to the ant-lion and other car-
nivorous creatures.
Rhadopterus picipes Oliv. [E. A. Schwarz]. One speci-
men of this beetle was taken July 30 from the jaws of
an ant-lion.
Coleopterous larvae. Some which could not be identi-
fied were often taken from the ant-lion pits. These had
evidently fallen from the vegetation above, and were
a considerable factor in maintaining so large a number
of thriving ant-lions.
Epicauta marginata Fab. [E. A. Schwarz}. About
August 20, several of these blister-beetles were found
dead on the bank. On September 4 and 6 there were
again large lots of these beetles, all dead. I cannot ex-
Plain the phenomenon, unless it be that some creature
had captured them and then spat them out, dead, be-
cause of their bad taste. :
Hyperodes humilis Gyll. [E. A. Schwarz]. One speci-
214 Trans. Acad. Sci. of St. Louis
men of this beetle was taken from the jaws of an ant-
lion larva on August 14, 1917.
Opatrinus notus Say. [E. A. Schwarz]. A dead beetle
taken from a spider’s web,
Harpalus dichrous Dejean. [E. A. Schwarz]. One dead
beetle found on top of the bank, September 4, 1917.
Anisotarsus sp. [E. A. Schwarz]. One individual of
this species visited the bank on September 6, 1917.
Sphenophorus parvulus. Gyll. [E. A. Schwarz]. This
snout-beetle was rescued alive from an ant-lion pit, where
the enemy was actively endeavoring to make a meal of
him on September 7, 1917.
Arhopalus fulminans Fab. [E. A. Schwarz]. Several
of these were found on the bank on May 28, 1917.
Languria mozardi Lat. [E.:A. Schwarz]. Three speci-
mens found on the bank on May 28, 1918.
Staphylinus cinnamopterus Grav. [E. A. Schwarz].
Of this rove-beetle, the only visitor was found on July
31, 1918.
Tanymecus confertus Gyll. [E. A. Schwarz]. This
snout-beetle visited the clay bank on May 20, 1920.
Orthoptera.
Orocharis saltator Uhl. [A. N. Caudell]. Six of these
were on a small plant growing at the base of the bank
on July 30, They probably contributed to the food sup-
ply of certain predacious inhabitants.
Gryllus pennsylvanicus Burm. [A. N. Caudell]. 4
small cricket was seen struggling in the jaws of an ant-
lion, into whose pit it had fallen on August 14, 1917,
and on August 30, one of this species spent a chilly night
in one of the bee burrows, and was the first to poke its
friendly head out into the light of day next morning.
The Ecology of a Sheltered Clay Bank 215
Ischnoptera pennsylvanica De Geer. An egg-case of
this roach was found in one of the old bee tunnels. Just
what would have been the chances of survival of the
young if they had hatched, we can only surmise. An
adult was also found hiding in a crevice on September 6.
Locust nymph. This visitor was found as prey of the
stick-bug, mesa brevipennis.
Nemobius carolinus Scud. [A. N. Caudell]. One dead
individual on the bank, September 7, 1917.
Melanoplus femur-rubrum. One resting here Septem-
ber 7, 1917.
Tettigidea lateralis var. polymorpha Burm. [A. N.
Caudell]. On October 3, 1917, several locusts of this
Species were on the clay bank, probably attracted to the
spot for shelter from the cold. One was in the web of
Steatida borealis Htz., where a battle occurred between
the plucky little spider and the big locust.
Hapithus agitator Uhler. [A. N. Caudell]. A male
visitor of this species fell prey to a spider, Phidippus
tripunctatus [J. H. Emerton] on September 6, 1919.
Hemiptera.
Pselliopus cinctus Fab. [H. E. Gibson]. This bug was
found in a spider’s web on September 12.
Anasa tristis De Geer. While this bug was seldom
seen at the bank during the summer, on October 3, a
dozen adults were found quietly at rest among the rub-
bish. Each year they appeared there late in the autumn.
These bugs hibernate as adults, and had come to
the bank seeking crevices in which to spend the winter.
They probably appeared upon the scene too late to in-
fluence the lives of others. On one occasion (May 20) a
young spider, Phidippus sp. [J. H. Emerton], was seen
feeding on one of these bugs.
216 Trans. Acad. Sci. of St. Louis
Reduvius personatus L. [A. T. McAtee]. This pro-
tectively colored bug, concealed in a crevice was sucking
the life-blood from a pioneer inhabitant, Ancistrocerus
fulvipes, on June 28, 1920, and earlier in these pages I
state that this bug was oS upon by the spider,
Ariadna bicolor.
Arachnida.
Phidippus. clarus Keyserling. [C. L. Shoemaker].
One specimen was found on the bank on July 16, 1917.
It was probably in quest of food. This is one of the
‘‘wandering’’ spiders, and its wanderings into the unit
were evidently not without purpose.
Phidippus sp. [J. H. Emerton]. A female actually
made a permanent home in one of the burrows, where
her face could often be seen peering from the doorway,
and occasionally she would catch and devour a Chalcid.
Phidippus tripunctatus [J. H. Emerton]. One young
one feeding upon a visiting male cricket Hopithus agt-
tator Uhler. [A. N. Caudell], on September 6, 1919.
Prosthesima sp. [J. H. Emerton]. A female was walk-
ing on the bank, June 28, 1920.
Thanatus lycosoides Emerton. [C. L. Shoemaker]. One
specimen found at the base of the bank on September
23, 1917.
Habrocestum pulex Htz. [J. H. Emerton]. One speci-
men on the bank September 23, 1917, feeding upon an
ant, Crematogaster lineolata.
Lycosa scutulata Htz. On July 30, 1917, at twilight,
a male of this spider was quietly resting at the
base of the bank. The Lycosidae are popularly know?
as wolf spiders; they are vagabond, hunting spiders,
spinning no snare but chasing their prey upon the
The Ecology of a Sheltered Clay Bank 217
ground, This sly watcher undoubtedly had business to
transact at the clay bank.
Lycosa sp. During the first part of August, several
were seen repeatedly at the same place. It seemed that
twilight brought out these night prowlers.
Xysticus ferox Htz, [C. L. Shoemaker]. One speci-
men at the clay bank at dusk, July 30. This is one of
the crab spiders, and like the others, probably came
in quest of food.
Spider. A small unidentified specimen taken from
the jaws of the ant-lion on September 3, 1917.
Pardosa sp. [J. H. Emerton]. This was another of
the wandering hunters which came to the bank October
3, when food elsewhere became scarce, and one specimen
was walking about on the bank one sunshiny morning
of February 26, 1921. :
Drassus sp. [J. H. Emerton]. One at bank October
3, 1917.
Gonyleptes sp. Several of these daddy-long-legs were
on the bank, and about a dozen had congregated on a
small plant on July 30, 1917. Warburton* says the
members of this group are ‘without doubt essentially
carnivorous.’? Hence the presence of these was prob-
ably of serious significance, especially to the smaller
Species of residents.
Ixodes sp. This tick was abundant each year at the
end of July and in the early part of August among the
old lumber on top of the bank.
Myriapoda.
On several occasions during 1917 and 1918, dead speci-
mens of Myriapods were found on the bank. Since none
*Cambridge Nat. History 4:44, 1909.
218 Trans. Acad. Sci. of St. Louis
were found alive and often the dead ones were partly
disintegrated, and since these species are known to
limit their habitation to rotten woods, I could think of
no explanation for their presence here in such condi-
tion except the far-fetched theory that they had been
brought here by other animals, possibly lizards, and
abandoned or spit up. ‘Among those thus found were
Euryeurus erythopygus Brandt. [R. V. Chamberlain],
Cleidogona caesioannulata Wood. [R. V. Chamberlain],
and Julius sp. [R. V. Chamberlain].
Résumé of Visitors.
The following table recounts species found among the
visitors to the clay-bank community:
Snake,
Lizards,
Toads,
Phoebe bird,
Red bird,
Ground mole,
Man,
Lepidoptera, Pyralis cuprealis.
Caterpillars, Pyralididae and Noctuidae.
Lepidoptera, Pseudaglossa lubricalus.
Catocala moth, Catocala innubens.
Moth, Herculia olinalis.
Wasp, Pompiloides sp.
Spider wasp, Arachnophoctonus ferrug’
neus.
Wasp, Pompiloides americanus.
Wasp, Tachytes peptictus.
Wasp, = Silaon sp.
The Ecology of a Sheltered Clay Bank 219
Pompilid wasp,
Leaf-cutter bee,
Bee,
Parasitic bee,
Beetle,
Lightning beetle,
Beetle,
Beetle,
Beetle larva,
Blister beetle,
Beetle,
Beetle,
Beetle,
Beetle,
Beetle,
Long-horned beetle,
Beetle,
Rove beetle,
Snout beetle,
Orthopteron,
Cricket,
Wild roach,
Cricket,
Locust,
Sphex nigricans.
Sphex procera.
Odynerus unifasciatus.
Polistes variatus.
Hoplisus phaleratus.
Trypoxylon nigrellum,
Miscophus americanus.
Vespa germanica.
Pseudagenia architecta.
Megachile generosa.
Halictus pectinatus.
Photopsis sp.
Camponotus herculeanus.
Corymbites inflatus.
Photinus pyralis. °
Hypera punctata,
Rhabdopterus picipes.
_ Unidentified.
Epicauta marginata.
Hyperodes humilis.
Opatrinus notus.
Harpalus dichrous.
Anisotarsus sp.
Sphenophorus parvulus.
Arhopalus fulminans.
Languria mozardi.
Staphylinus cinnamopterus.
Tanymecus confertus.
Orocharis saltator.
Gryllus pennsylvanicus.
Ischnoptera pennsylvanica.
Nemobius carolinus.
Melanoplus femur-rubrum.
220 Trans. Acad. Sci. of St. Louts
Orthopteron, Tettigidea laterals var.
polymor pha.
Orthopteron, Hapithus agitator.
ug, Pselliopus cinctus.
Squash bug, Anasa tristis.
Bug, Reduvius personatus.
Spider, Phidippus clarus.
Spider, Phidippus tripunctatus.
Spider, Phidippus sp.
Spider, Prosthesima sp.
Spider, Thanatus lycosoides.
Spider, Habrocestum pulex.
Spider, Lycosa scutulata.
Spider, Lycosa sp.
Spider, Xysticus ferox.
Spider, Pardosa sp.
Spider, Drassus sp.
Daddy-long-legs, Gonyleptes sp.
Tick, Ixodes sp.
Myriapod, Euryeurus erythopygus.
Myriapod, Cleidogona caesioannulata.
Myriapod, Julus sp.
The list totals 71 species, and covers a ionhe of
forms, from man down to Myriapoda. No attempt was
made to list the microscopic organisms, although with
proper equipment an interesting list of such specimens
could doubtless have been found.
It is interesting to note that of these 71 species which
were found visiting the clay bank coincident with my
intermittent visits 7 were vetebrates, 5 were species of
Lepidoptera, 18 Hymenoptera, almost all of which were
wasps, 15 were beetles, 7 Orthoptera, 3 species of bugs,
10 Arachnida and 3 Myriapoda.
The Ecology of a Sheltered Clay Bank 221
It will be seen that some of the creatures were of
little or no importance in the balance of life on the clay
bank; for instance the squash-bug, Anasa tristis, which
is a vegetable feeder and which few creatures would
eat, would have little influence on the life of the social
group. On the other hand, some of the visitors wielded
a significant influence, as in the case of Vespa germanica
which carried off the Chalcid parasites which preyed
upon the bees. Many of the visitors might be placed
alternatively in still another group, those which might
properly be considered as either renters or pioneers,
as for instance, the spider-hunter, Pseudagenia archi-
tecta, which might be said to carry on its daily busi-
ness at the bank, although it maintained its home else-
where. Of course the sharp line of demarcation between
the groups is difficult to define while the possibility is
always open for the visitors to become residents there
and participate in all the joys and sorrows incidental
to influencing the balance of nature already present,
by preying, by being preyed upon or by parasitizing
other creatures or by becoming hosts for other para-
Sites,
(e) Parasites.
The following account presents the details of the
activities of the last group of the inhabitants of the
bank, the parasites, which affected the pioneers and the
renters. These exerted their influence upon the balance
of population in a variety of ways, and in turn were 1n-
fluenced in interesting ways by some creatures from
all three groups.
222 Trans. Acad. Sci. of St. Louts
Parasitic Hymenoptera.
Chalcid parasites, Monodontomerus sp. [J. C. Crawford].
When the observations began on June 25, 1917, and
the Anthophora bees, etc., first appeared that spring,
hundreds of these parasites were at once on hand. Their
emergence was almost simultaneous. The number on
this first day of my visit was so great as to make me
fear the appalling mortality that must be in store for
their hosts, the bees, even if only a small portion of
them were to be permitted to beget offspring. They
were actively searching over the bank, entering burrows
and dancing before the bank in the bright sunshine.
I could then get direct evidence only that they had come
from the cocoons of Anthophora abrupta, but they were
undoubtedly parasitic upon other species as well, since
they were found at the bank also in the fall, and the
bees, A. abrupta, ended their life cycle in July. One
burrow of the mining-bee, when opened, exposed to
view fifty Monodontomerus adults; I could not be ab-
solutely sure, however, that all of these were young
parasites just emerging from this cell; it is possible
that some of the sun-dancers had left their companions
and crowded into this cell of mixed sexes. In 1920, on
July 24, 13 cells of this Anthophora bee were taken into
the laboratory, only four of which proved to be infected
by this parasite. These four cells gave forth 139 adult
parasites, 109 of which were females and 30 males, or
an average of about 35 for each bee cell. At such an
alarming rate of increase, it is surprising that any bees
at all should have escaped them.
These parasites throve and appeared in large num
bers in each of the seasons of observation. There 1#
The Ecology of a Sheltered Clay Bank 223
more than one generation each year. This made a par-
ticularly difficult situation for the species of hosts which
produced but one generation each year. The first adults
that emerged in the spring, lived and disappeared
simultaneously with the Anthophora that closed their
life cycle about the middle of July. The eggs deposited
by the first lot of adult parasites produced adults later
in the summer. A careful and frequent examination
showed that the Chalcids were not in evidence from the
time of their waning and first disappearance, July 16,
up until September 3, when they re-appeared after an
interval of forty-eight days. This second generation
also comprised an enormous population. Many of these
were seen also to enter the bee burrows, and some of
the bee cells dug up later also contained these Chalcids
in the pupal stage.
Not all of them succeeded in their parasitic plan of
existence; some fell prey to other inhabitants early in
life. On the first day of the appearance of the second
brood, seven were taken dead from the spiders’ webs
nearby, and elsewhere in these pages is recorded the
fact that Vespa germanica was making regular trips,
carrying them off day after day to her nest.
On September 6, while the numbers flying about seemed
just as great, still many were found dead about the bank;
[ cannot account for this, since this early date seems
hardly time for a natural death. These may have been
the dead of the previous generation, or those of the
Present generation that had perished before emergence,
Which some industrious Trypoxylon wasps had swept
out of the old burrows that they were renovating for
their own use. Mortality was really upon them, how-
ever, because by September 12 a marked reduction in
224 Trans. Acad. Sci. of St. Louis
their numbers was easily noticeable. On October 3,
none at all were abroad, but of course at this date prac-
tically all of the life on the bank was still.
In 1918, when the carpenter-bees, the Anthophora
mining-bees, and the caterpillar wasps all appeared a
month earlier, about May 25 (because of temperature
conditions), these Chalcid parasites too appeared a
month earlier. All of the life in the bank responded to
the same laws. In 1918 these parasites were just as
numerous but I was surprised to note no increase over
the previous year. It would have seemed reasonable
to expect that the myriads of the last year would have
multiplied as greatly, at the expense of everything else,
that this year there would be little or nothing but Chal-
cids. But to our surprise the 4. abrupta bees, even under
such persecution, were more abundant the second year
than the first. This is a good example of the delicate
balance in nature’s game of cat and mouse.
By June 28, 1918, as many chalcids were dead and
strewn about the bank, in burrows and in crevices, a8
were alive and on the wing; thus that date marked the
decline of the first generation, which in the preceding
year occurred on July 15. SomeAnthophora cells taken
a little latter contained live pupae. So, comparing the
dates of this species for 1917 and 1918, we see that when
the first generation got a thirty days’ earlier start, 80
likewise did the second generation.
In 1920 this species appeared in about the same num-
bers, and the interval dividing the two generations again
came in July. To be exact, on my visit on July 21, not
one Chaleid was to be scen. But on the next inspection,
September 2, they were out again in great numbers.
These of the second generation were, in turn, all gone
The Ecology of a Sheltered Clay Bank 225
the first of October. The dates of these two generations
were practically the same as in the first year; the first
generation ran from late June to mid-July, and the see-
ond appeared about the first of September but disap-
peared before the end of the month,
The animal most abundant in an ecological unit
should be regarded as the dominant one. If this rule
includes parasites, then we must admit that this Chaleid
is the most abundant and dominant insect in the unit.
They were already plentiful at the time of my first ob-
servation; their hosts were well established and numer-
ous, and since each host is capable of bringing to matur-
ity, on an average, thirty or so parasites, we see how fa-
vorable were all their prospects. And yet we come face
to face with the fact that other parasites, Diptera, etc.,
even though they appear in far smaller numbers, di
fully as much destruction to the permanent residents of
the bank as did these many little Chalcids. What does
it matter to an Anthophora larva, whether it yields its
life to produce thirty-nine chalcids or one cuckoo-bee?
Thus, we see that the most important aspect of the
life of the Chalcids is to keep the bees from overrunning
everything by their enormous increase in population,
which would doubtless take place without this deterring
influence. As a secondary function they serve as food
for some species that in no other wise influence the life
of the bank, such as non-resident Vespa germanica, and
also to feed spiders which, by being thus encouraged
to come to or stay on the bank, might also prey upon the
other residents.
226 Trans. Acad. Sci. of St. Louis
Cuckoo-bees and parasitic wasps.
These beautiful green bees, parasitic upon many
forms of Hymenoptera, appeared about the bank dur-
ing every season of the term of observation. They were
directly parasitic upon many of the inhabitants, and
were frequently observed following a returning mother
to her nest, and were often seen prowling about and
entering the nests of others.
Only one species was found, Chrysis (Tetrachrysis)
coeruleus Fab. [S. A. Rohwer]. These cuckoo-bees ap-
peared with the first life on June 25, 1917, and were
in almost constant attendance at the bank in summer
from that date until near the middle of October. They
appeared in great abundance in early summer, flying
before the bank in the sunshine, searching over the sur-
face and entering numerous crevices and bees’ tunnels,
evidently taxing the legitimate life in the bank as much
as did the Chalcids. Their numbers seemed to increase
as summer advanced; but numerical estimates of this
population are difficult to make at the early season. One
may count the turrets of the bees, the pits of the ant-
lions, or the webs of the spiders, but these transients
sneaking about the dwellings of others, could not be
definitely enumerated.
The next year by August 10 their number seemed
much reduced; on August 14 only one specimen Was
seen, and none thereafter, excepting dead ones at the
foot of the bank. One carcass of this bee was removed
from the web of a spider. This was probably the
natural time for the death of these bees, and we thought
that a new generation would appear soon after. It was
interesting to discover that with the disappearance of
The Ecology of a Sheltered Clay Bank 227
the cuckoo-bees, the silver-winged parasitic Dipteron
made its appearance. '
On August 30, only one cuckoo-bee was present; this
seemed a forerunner of the second generation, rather
than a straggler from the first, for the next day three
were out, and thereafter they increased briskly until a
thriving second generation was out and in action. From
September 4 to 6 this population was at its height; by
September 12 they, along with the other species then
about the bank, began to disappear as the chill nights
came on, but whether they perished with the cold, or
made their natural demise, or merely were in hiding, I
cannot say with certainty. However, on one late Sep-
tember day, when the bank became warm in the sun-
shine, two appeared; one was especially active in annoy-
ing a turret-bee. No doubt many of these parasitic bees
died in their burrows. Their latest date recorded was
October 13.
Few of these superchitinous insects fall prey to the
hungry neighbors. The rule ‘‘to eat and be eaten’’
does not apply to the cuckoo-bee, which appears to be
the armadillo of the insect world.
Pseudomelecta interrupta Cress, [S. A. Rohwer]. This
bee was seen only once at the clay bank, on July
21, 1920. It entered one of the nests of Anthophora,
and was taken for identification as it emerged several
minutes later. hice
The species must have a highly interesting life his-
tory, for Wheeler (Proc. Am. Phil. Soc. 58:14. 1919)
classifies it as one of the parasitic bees, derived from
the ancestral genus Anthophora and probably parasitic
on Anthophora. He questions this later statement, but
since it entered the burrow of Anthophora after the
228 Trans. Acad. Sci. of St. Louis
manner of a parasite, I think much doubt as to its habits
is removed. This of course would mean that P. inter-
rupta, having at some time been derived from the genus
Anthophora has found a way to escape the Anthophora
responsibilities of mining and food gathering by turn-
ing parasitic on its own kin, just as seems to be the
case with Psithyrus in the nest of Bombus.
Mutillids. The cow-killers or velvet ants, with wing-
less females and winged males, are well known to be
parasitic, and their way of prowling about the clay bank
indicates that, like many of our own species, seeking
hosts is the sole ambition of their lives. They undoubt-
edly found hosts in the clay bank.
Dasymutilla ferrugata Fab. [S. A. Rohwer]. On June
28, 1917, one female was seen to enter a dozen
burrows of the Anthophora bee, and in some it remained
a sufficiently long time to have done mischief. Other
mutillid parasites, which remained unidentified, were
seen doing precisely the same on July 31, and August 10.
Sphaerophthalma scaeva. Two females of this para-
site were darting about as if on mischief bent, on Au-
gust 30, and on the next day a half-dozen or more were
seen. We have previously reared these parasites from
the old nests of the mud-dauber, Sceliphron caementar-
qum.,
On the clay bank, besides D. ferrugata and S. scaeva,
one would occasionally pick up an unknown specimen of
the Mutillids; I suspect that they were seeking hosts,
but their numbers were so insignificant in comparison
to the other parasites that I do not think they were to
be regarded as a ‘‘real force in the community.”’
Sphaerophthalma pennsylvanica Lep. [S. A. Rob-
wer]. A small elm shrub, between the clay bank and
The Ecology of a Sheltered Clay Bank 229
the rose bushes, seemed especially attractive to a large
variety of insects. Flies and parasitic Hymenoptera
were particularly abundant in variety, but for large
numbers of a single species only two were notable; they
were Chalybion caerulewm males, discussed elsewhere
in these pages, and males of this species. There was
another similar shrub near by, but no such insect popu-
lation was apparent upon it, so I concluded that it was
the aphids present here which had attracted them to
this one, although I could see no actual relationship ex-
pressed in their activities. Perhaps in both species,
the males were merely marking time until the appear-
ance of the females. It is interesting to note, how-
ever, that while the males were so near to the clay bank
on this occasion, they probably flew here from else-
where, for the females of this parasite have never been
seen at the bank.
Sapyga sp. [S. A. Rohwer]. One was walking about
on the bank on June 29, 1917. Members of this family
are, according to Rohwer (Conn. Nat. Hist. Survey
22:620. 1916), parasitic on bees and wasps, while
Packard (Guide to Study of Insects, p. 134. 1876) says
that in Southern Europe Sapyga repanda is parasitic
in the nests of Xylocopa violacea; Sharp, however, (In-
sects, Pt. 2, p. 100) says that Sapyga 5— punctata was
seen carrying caterpillars. :
Sphecodes sp. Two specimens of this red-bellied
parasitic bee were seen going in and out of the burrows
of the resident bees on June 28, 1920.
Diptera.
Of all the Diptera taken at the bank, not one speci-
men was other than parasitic; the group as a whole may
230 Trans. Acad. Sci. of St. Louis
be justly regarded as doing as much damage to the law-
ful inhabitants as either the Chrysits or the Chalcids.
The Diptera, however, came later in the season, and
possibly sought different victims. Dipterous parasites
were first seen on July 30, 1917—a half-dozen represent-
ing two or three species. As days went by, they became
more numerous in species and numbers. The details be-
low relate their activities.
Copecrypta ruficauda Coq. [C. H. T. Townsend]. This
fly was seen to rest for several minutes on the clay
bank on August 10. While I was wondering how I
could capture it, it flew to my wrist and became so in-
tent on drinking the perspiration that I picked it up
easily in my hand. It was the only specimen seen there.
Archytas aterrima R. D. [C. H. T. Townsend]. This
fly belonging to the family Tachinidae, and commonly
called the tachina fly, is parasitic on other insects. One
specimen was taken at the bank on July 31. Like Cope-
crypta ruficauda, this fly was taken when it flew to my
hand to drink the perspiration.
Sarcomacronychia trivittata T. [C. H. T. Townsend].
The first time this fly was seen at the bank was
September 6, when two specimens were taken. They
spent their time flying about the bank, often resting
upon it and seldom going far away from it; this con-
duct indicated that there must be some attraction there.
A third individual was observed on October 3, 1917.
Parametopia sp. [C. H. T. Townsend]. One such pest
was seen to follow the wasp, Ancistrocerus fulvipes,
as she entered her nest laden with a caterpillar on Sep-
tember 12. A second individual was scen to follow the
same species of wasp on October 3, 1917, as she was 8%
ing to the nest. This does not mean that this fly has
The Ecology of a Sheltered Clay Bank 231
become so specialized as to require a specific host, for
we have often seen it behave so with other wasps else-
- where,
Argyromoeba anale Say. One specimen of this para-
sitic fly was on the bank September 7, 1917.
Argyromoeba oedipus Fab. [F. Knab]. Six specimens
of this parasite were seen about the clay bank on July
31, one of which actually entered a burrow of the white-
banded bee, Entechnia taurea.
Argyromocba fur O. S. [F. Knab]. One was at rest
on the clay bank on September 3, 1917. In 1918, this
species, along with all the other life in the unit, appeared
about a month earlier, or on May 28. (The 1918 material
was identified by C. T. Greene.)
Argyromoeba tigrina De G. [F. Knab]. This para-
Sitic fly (Fig. 18) first appeared on August 8, 1917. At
first there were only three individuals; two days later
they were present in very great numbers. They frisked
and flitted about in the sunlight, stopping to rest for
long periods on the wooden part of the porch and in the
direct sunlight. When at rest thus they could easily be
Picked up with the fingers. The peculiar life history of
this fly enables us to know absolutely that they had not
flown to the bank from afar, but had actually emerged
on the scene. The tell-tale evidence is this: in trans-
forming to adulthood, this species leaves the home of
its host while it is still in the pupal stage and, upon
gaining its freedom transforms into the adult and
leaves the old shedding-skin (Fig. 19) on the surface
near the point where it emerged. In many places on
the face of the clay bank, and at its base, up among the
Wooden partitions very near to the tunnels of the car-
Penter-bees, Monobia wasps or grass-carrier wasps, and
232 Trans. Acad. Sci. of St. Louis
even protruding from the bee burrows, were these shed-
ding skins adhering conspicuously. This indicated just
which species were probably their hosts. That they
were quite numerous I was also certain, since I picked
up more than sixty of these shedding-skins, and of course
these were not all.
While there is sufficient evidence that these para-
sites prey upon the inhabitants, and while they spend
most of the sunshiny hours each day in flying to and
fro before the openings of the burrows or resting near
by, and although I have watched for hours at a time, I
have never yet seen one of these creatures attempting
to enter a burrow. I especially watched for this, since
the great Fabre has discovered for another species of
this genus some complicated aspects of life history.
True I have often seen the females hovering over a bur-
row, poising on the wing above the aperture and dipping
down again and again, much in the manner of a dragon-
fly depositing her eggs under the water, and I suspected
that with each dip she deposited an egg but, like Fabre,
I have never discovered the egg. Fabre seems to think,
for his species, that the parent fly oviposits by merely
dropping a minute egg while flying over the surface of
the mud walls which contain the grubs of the host, and
that the larvae hatching from the eggs are wonderfully
adapted for breaking into the masonry to reach the host
by being provided with a very horny, deflexed head,
armed in front with stiff bristles and under the body with
several pairs of elongate setae serving as organs of
locomotion. In the species under present consideration,
the eggs are probably deposited in this manner, although
some other species of this genus, as A. oedipus, ate
known to enter the bee burrows. Several females of A.
The Ecology of a Sheltered Clay Bank 233
tigrina were seen repeatedly to fly violently against the
wood beside the Xylocopa or other burrows; then often
I could see the tip of the abdomen enlarge, as it was
bent downward and tightly pressed against the rough
surface, and moved in a circular motion as it rubbed
against the wood. In only one instance did I actually
See something fall, but I was not near enough to see
whether it was an egg or excrement. So the problem
of the life history of this parasite remains unsolved.
It would at first appear that A. tigrina came too late
in the season to affect the bees, Anthophora abrupta,
since their lives had terminated before the advent of
the parasites. This might be true, and A. tigrina might
not affect this species but might be a direct cause of
the gradual lessening of the numbers of the white-
banded bees, Entechnia taurea, which were contempo-
raneous. However, if Fabre’s theory holds true for this
species and these parasitic young possess those won-
derful adaptations for reaching their host even through
prison walls, then the parasites would affect the dormant
generation of Anthophora, and other species as well.
There is additional evidence, however, in the fact that
I found the shedding-skins on the face of the bank, but
not on the top, and A. abrupta builds only in the face of
the bank. This would lead us to think that A. abrupta
is a host despite the fact that the adults are not con-
temporaneous.
I want to repeat that in constant watching during
these dipping maneuvers, I failed to see an egg drop, but
from the insect’s behavior I felt sure that something was
happening. It is reasonable to believe that the egg is
minute, and with the observer four feet away (one can-
234 Trans. Acad. Sci. of St. Louis
not successfully come closer) something might happen
beyond one’s sense of sight.
These adults are flower-loving insects. On August 13,
two specimens were seen on flowers in a field some dis-
tance away. It may well be that this would account for
the reduced numbers during the morning hours, while at
noon they were present in full force; perhaps they were
out in the blossoms satisfying their morning hunger, or
again perhaps they had gone out into the vegetation to
spend the night (their sleeping habits have not yet been
ascertained) and were slow in returning to the day’s
activity. With the cold nights of early September they
began noticeably to wane, and none were seen alive after
September 12.
These flies were seen throughout the season in vary-
ing abundance. It seemed at first that with their play-
ing and dancing about the bank they were merely mark-
ing time, but, since they continued it all summer, it ap-
pears that they are the kind of beings that can seemingly
dance and frolic through life and yet make a success of
living. Some were seen in copulo on July 26, 1922.
One pair remained in copulo, back to back, for a half
hour, resting on a shady portion of the bank.
In 1918, this fly was no exception in the phenomenot
of all life appearing a month earlier. A few individuals
were seen out on July 17, 1918, as against ‘August 1,
1917. Until the end of July they were very abundant,
and like their ancestors of yesteryear, they kept busy
most of the time, doing nothing in particular, unless
they oviposited on the sly. The shedding-skins as well
as the flies gave evidence of an increase in population.
This increase was probably at the expense of the dwellers
in the wood-tunnels above, for as I have already show?
The Ecology of a Sheltered Clay Bank 235
these and the white banded bees declined simultaneously
with the increase in this parasite.
I do not know what happened to the species in 1919,
but in 1920 they appeared in limited numbers, and in
1921, they were again on the increase.
This fly evidently has a sensational criminal life his-
tory, which I hope some day will be brought to light.
Ganperdia apivora Aldrich [J. M. Aldrich]. In June,
1920, a dozen or more specimens of this red-eyed diptron
were flying about the burrows in the clay bank, in a
manner strongly indicating parasitic habits, although I
did not at any time actually see them enter the burrows,
and the same observation was made on May 27 to May
28, 1921. These bees are undoubtedly parasitic on the
mining bees in the clay bank, since I actually observed
them hatching in the laboratory from cells of Antho-
phora abrupta on May 20 to 22, 1921, from material
taken about nine miles north of the clay bank.
This species is described by J. M. Aldrich* from spec-
imens taken near St. Louis in 1877. It is very interest-
ing, too, that forty-five years ago this fly was found in
a clay bank about the nests of the mining bee, Antho-
phora abrupta. That it is parasitic upon these bees is
evident from the statement in the paper cited, that one
‘of the bee cells contained cocoons which gave forth
adult flies of this species in the following Spring.
In the first week of June, 1922, hundreds of these red-
eyed flies were out at the bank and were then being
freely preyed upon and devoured by the lizard (Cnemt-
dophorus sealineatus) which made itself at home about
the place.
*Proc. Ent. Soc. Wash. 21:106. 1919.
236 Trans. Acad. Sct. of St. Louis
Lepidophora lepidocera Weid. [C. T. Greene].
This parasite was present at the clay bank on July 1,
1922. It was often seen bobbing up and down on the
wing before the burrows, but if eggs were dropped in
the process, none were ever discovered.
Parasitic Beetles.
Hornia minutipennis Riley [E. ‘A. Schwarz]. This
beetle, which is parasitic upon the mining bees, was
found only twice at the clay bank. One was taken at the
base of the bank on May 27, 1920; it deposited many
eggs in captivity the next day, and died soon after. An-
other was taken from the cell of A. abrupta on October
2, 1920; it emerged in the warm laboratory on March
19, 1921, promptly ate of a piece of apple and consumed
part of an Anthophora larva which was given it; the re-
mainder of the larva it allowed to dry up. It lived five
weeks. 7
C. V. Riley* describes this species from specimens
reared from the cells of Anthophora sponsa taken from
elay banks near St. Louis. Piercet tells us that the
genus Hornia is known to be parasitic on Hymenoptera;
the larvae are conveyed to the host by some other insect
—that is, they are passively conveyed. Their feet are
specialized for clinging, not digging. Their food is
generally honey; hence the mouth parts are reduced.
Hornia shows degeneracy in the adult stage with com-
plete loss of the wings, and almost entire reduction of
the elytra.
Fabret has found a parasitic beetle, Sitaris humeralis
*Trans. Ent. Soc. St. Louis, 3:563-565. 1877.
Univ. Nebraska Studies, 4:153. 1904.
arp, Insects. Pt. II, p. 272.
The Ecology of a Sheltered Clay Bank 237
by name, living at the expense of bees of the genus
Anthophora. ‘‘The eggs of Sitaris are deposited in the
earth in close proximity to the entrances of the bees’
a ar a single female producing upwards of two
thousand eggs. ..... They hatch in about a month,
producing a tiny triungulin of black color; the larvae,
however, do not move away, but, without taking any
food, hibernate in a heap until spring, when they be-
come active. Although they are close to the abodes of
the bees, they do not enter them, but attach themselves
to any hairy object that may come near them, and thus
get onto the bodies of Anthophora and are carried to the
nest.’’? At first it seemed strange that at the clay bank
at Wickes they were present only in such small num-
bers, but this is explained by the statement from the
Same work that the triungulins distribute themselves
on all sorts of unsuitable insects, so that it is possible
that not more than one in a thousand succeeds in gain-
ing access to the Anthophora nests.
Résumé of Parasites.
To summarize this group, I have found in the clay
bank parasites to the number of 19 species. This
is highly interesting in the light of the relation of
this number to the number of pioneers and renters.
Of the first there were 18 species, many of which could
not possibly be host of these parasites; of the latter
there were 11 species, including spiders and others which
likewise could not serve as host. Leaving out of consid-
eration all the non-host species of both the pioneers and
the renters, we find the parasites greatly outnumbering
legitimate residents of either class.
The following table recounts the parasites that visited
the bank.
238 Trans. Acad. Sci. of St. Lowis
3 Parasites.
Chalcid parasite, Monodontomerus sp.
Cuckoo-bee, Chrysis (Tetrachrysis)
coeruleus.
Parasitic bee, Pseudomelecta interrwpta.
Cow-killer or velvet ant, Dasymutilla ferrugata.
Cow-killer or velvet ant, Sphaerophthalma scaeva.
Cow-killer or velvet ant, Sphaerophthalma pennsyl-
vanica.
Parasitic Wasp, Sapyga sp.
Parasitic Bee, Sphecodes sp.
Parasitic Fly, Copecrypta ruficauda.
Parasitic Fly, Archytas aterrima.
Parasitic Fly, Sarcomacronychia trivtt-
tata.
Parasitic Fly, Parametopia sp.
Parasitic Fly, Argyromoeba anale,
Parasitic Fly, Argyromoeba oedipus.
Parasitic Fly, Argyromoeba fur.
Parasitic Fly. Argyromoeba tigrina.
Parasitic Fly. Ganperdia apivora.
Parasitic Fly. Lepidophora lemdocera.
Parasitic beetle. Hornia minutipennis.
It will be seen from the table that, with the exception
of one beetle, all of the parasites belonged to the orders
Diptera and Hymenoptera. Of the former there were
10; of the latter, there were 8, comprising one chalcid,
three species of cow-killers or velvet ants, one wasp and
two species of bee which only recently phylogenetically
speaking, have acquired parasitic habits, and one spe-
cies of cuckoo-bee. With this appalling array it is @
wonder that any host can continue to exist; there is 20
The Ecology of a Sheltered Clay Bank 239
telling how many species have already become extinct
through this agency during the existence of the clay
bank.
(C) RELATION OF POPULATION TO ENVIRONMENT,
(a) Relation to Temperature.
Insects know not dates; May or December, June or
January mean nothing to them other than heat or cold.
The insects’ calendar is the thermometer. Instead of
saying that all the spring Hymenoptera, regardless of
species, emerged during the latter part of June, I should
say that they emerged when the average mean tempera-
ture for fifteen days reached 73°. Let us not forget that
June has nothing in its makeup to mean anything to the
Hymenoptera, but 73° for fifteen days at the springtime
of the year means to them life, sunshine and food.
My attention was first attracted to this phenomenon
in noting that in 1918 the life appeared from the clay
bank about a month earlier than in 1917, and in 1920 ©
they appeared at almost the same time as in 1918. In
other words, the hymenopterous life about the place
was in its spring fullness on June 25, 1917, May 28, 1918,
and May 30, 1920. The average mean temperature for
the fifteen days preceding and including June 25, 1917,
was 73 1/25°; the average mean temperature for fif-
teen days preceding and including May 28, 1918, was
73 4/25°; and for the same period preceding May 30,
1920, was 731/2°. These temperature figures of course
are only relative; the clay bank was facing the eastern
sun, whereas, the temperature records were made in the
shade; let us remember, too, that the life potential buried
in the clay bank was likewise in the shade.* This corre-
*The temperature records are from the U. S. Meteorological Re
ports taken at St. Louis, twenty miles north of this site.
240 Trans. Acad. Sct. of St. Louis
lation works out so beautifully that I shall want to place
this material experimentally at a temperature of 73°
in November or February, to see if then the develop-
ment and maturity occurs likewise. Had the 1917 pop-
ulation (which emerged in abundance about June 25th)
emerged on May 28, as did the 1918 lot, it would have
spent the 15 days before its emergence at a temperature
of 66 2/15°.
This temperature phenomenon seemed to hold in gen-
eral for most of the spring Hymenoptera, and I found
them all appearing at one time, after the manner of
Anthophora abrupta for example. The bank yielded
its contents of this species all at one time each year,
within a space of two or at most three days. Simulta-
neously appeared also their Chalcid parasites, Mono-
dontomerus. The green cuckoo-bees burst forth at their
appointed time, and so too the caterpillar wasp, Ancis-
trocerus fulvipes; the Monobia mud wasp, the Trypoxy-
lon, ete., ete. The only exception was the carpenter-
bee, Xylocopa virginica which appeared some time
earlier.*
This species is a wood dweller—not a cave dweller—
and would, therefore, be influenced by temperature
somewhat differently from those which live in the com-
pact earth. Not alone in 1917 did I see this simultane-
ous emergence of the various species and of all individ-
uals of each of these species, but in 1918 and 1920 as
well. It was as though some one touched an electric
button and that button was labeled 73° F.
The white-banded bees E. taurea and the dipterous
parasites, demanding a higher temperature and hence
*We do not know how this bee hibernates, but observations
given elsewhere indicate a period of hibernation in the adult stage.
The Ecology of a Sheltered Clay Bank 241
emerging later, conformed nicely to the precedent set
by the spring Hymenoptera in the year 1918, when the
meteorological conditions were very different from 1917.
This bee population emerged in 1917 on July 16 and in
1918 with a more favorable temperature 18 days earlier
on June 28. In this species I found, in so far as the
records were kept, that they responded, by emerging
from the pupal condition, to a temperature of 77°. Using
the same weather records as a basis, we find that for the
fifteen days preceding their emergence on July 16, 1917,
we had an average mean temperature of 77 2/15°. In
1918, their appearance was earlier, June 28, and the
average mean temperature for the period of fifteen days
preceding this was 77 2/3° F.
Thus, we can see in the emergence of this hymenop-
terous population a very definite response to tempera-
ture, and not an emergence correlated with the length of
time the organisms had spent in the ground as larvae or
pupae. This is quite reasonable, as is already known,
the time spent in the various stages is lengthened by
cold, and the development is accelerated by warmth.
‘All this seems, in so far as these Hymenoptera are con-
cerned, not to apply merely to a single species, but to
indicate a law influencing all life under these conditions.
(b) Relation of population to light and sunshine.
The Hymenoptera are generally regarded as lovers of
the light and sunshine, and especially is this true of
the flower-frequenting bees and wasps; they are gen-
erally to be seen foraging among the blossoms, and then
indeed may we call them children of the sun. Here we
are interested in the three phases: (a) the relation of
the position of the nests to the sunlight, (b) the rela-
242, Trans. Acad. Sci. of St. Louis
tion of the sun-loving parasites to the position of the
nests of their hosts, and (c) the action of parasite to-
ward host as the nest of the host is in the sunny or the.
shady portion of the bank.
Position of the clay bank in relation to the sunlight.
We have seen in Fig. 2, how the clay bank was
protected above by the porch and faced the eastern sun.
Three clumps of rambler rose-bushes stood in front of
the bank, one at the north end and two spread over the
south half. These were densely covered with flowers
and foliage during the summer.* Since two rose-bushes
were in the southern half, the portion of the clay bank
just behind them was much more shaded than the north-
ern half.
In this brilliantly illuminated area between the bushes
the nests of 4. abrupta abounded; here they left their
old burrows, an unmistakable record of the life which
had been there. It was in this portion of the bank also
that in 1917, the most of the contemporaries of the
Anthophora congregated to build their nests. It would
be better to say that here they emerged, and then re- —
mained in almost the precise spot to nidify, but this in
turn only means that their ancestors at some time ac
tually chose this spot or congregated here, so the signi-
ficance is the same. In order to realize the sharp con-
trast in the nesting activities, between the sunny poT-
tion of the bank and the shaded areas, Figs. 3 and 4
may be compared. Fig. 4 shows the northern half
(the photograph was taken after a dry year, when few
bees made turrets, and these few had fallen or had been
*This picture was taken in October, when most of the foliage
had fallen.
The Ecology of a Sheltered Clay Bank 243
harvested for study), or that portion which got the max-
imum amount of sunlight. This is in striking contrast
to Fig. 3, the southern half, which enjoyed only a lit-
tle filtered sunlight. One can readily see that the de-
gree of illumination made a great difference in choice of
site by the A. abrupta. The white-banded bees, En-
techma taurea, the caterpillar-wasps, Ancistrocerus ful-
vipes, and the Trypoxylon wasps did not give evidence
of so direct a dependence upon this factor. I have often
wondered just what is the correct explanation of this
correlation of warmth to longevity. I have discovered
in other studies that in organisms that take no food as
adults (Saturniid moths) increased warmth tends to
accelerate the activity and thus exhaust the vital energy
so that the life of the organism is cut to a much shorter
duration than it would have been at a lower temperature
where activity is retarded. So shall we say that be-
cause A. abrupta chooses the warmest place in the sun,
her activity is intensified and her life soon spent? Or,
on the contrary, shall we say that her habit of choosing
the sunny situations is a fortunate adaptation in that it
enables her to get through with the necessary duties of
life in the short period of time allotted to her, and that
probably only those individuals which do exercise this
choice will be able to finish their life-work and leave
progeny? It should be noted that all the other insects
mentioned in this study have a much longer time for
their duties than A. abrupta, since they either lived the
entire season, or had more than one generation each
season.
In 1917, the white-banded bees greatly outnumbered
the Anthophora. While a few of the nests of the former
were in the face of the bank, and often took a somewhat
244 Trans. Acad. Sci. of St. Louis
vertical position (Fig. 20E) most of them were hort-
zontally resting on top of the bank, near the edge (Fig.
21E). All of the nests of this species enjoyed an abun-
dance of the sunlight; those on top of the bank were
made very near to the front edge. However, during
the years that followed this species decreased in num-
bers so that by 1920 it was barely holding its own,
whereas A. abrupta increased amazingly, as described
elsewhere. Another interesting factor enters into the
story here, however; during the years under observa-
tion the white-banded bees gradually extended their nest-
ing area. In 1917 most of them were in among the An-
thophora nests near the sunlight, either in the face of
the bank or near the edge of the top (Figs. 20E and 21K)
with merely the beginning of a settlement at the extreme
south end and another at the north end. In 1918, they
were almost all in these two end groups, and none among
the Anthophora nests. In 1919 a few were at the south
end, but most of them were on the top, farther back than
previously, and in 1920 to 1922, all of the burrows found
were far back on the bank and entirely out of the direct
sunlight. In the meantime Anthophora abrupta was in-
creasing in numbers, always building on the sunny face
of the bank. Moreover they were gradually spreading
over the southern portion of the bank, until by the end
of the period the north half of the dark barren portion
seen in Fig. 3 was abundantly inhabited; Fig. 22 is
the same area as in Fig. 3. Not without cause, however;
was this significant change; in the spring of 1918 the
tenants of the house had cut the ramblers almost to the
ground, so for the seasons immediately succeeding, that
portion of the bank was also fully exposed to the
sunlight.
The Ecoloyy of a Sheltered Clay Bank 245
The parasites also love the sunlight. This point leads
us at once into a maze of interrelations between sun-
light, parasites and hosts. The very short life cycle of
Anthophora abrupta eliminates from consideration prac-
tically all of the Dipterous parasites, because by their
early emergence they escape the parasites and their suc-
ceeding generation is practically provided for before the
most of the parasites are born. Hence Anthophora has
to struggle against only the Chalcids, the cuckoo-bees
and possibly one species of Diptera in order to main-
tain her species. While these three parasites did evi-
dently prey upon this Anthophora population to some
extent, their success depended to a far greater extent
upon the white-banded bees as hosts since the duration
of the life of the latter was more than twice that of the
Anthophora bees, and since the white-banded bees emerg-
ing later in the summer were the possible hosts of more
than three times as many species of parasites. It is
little wonder then, that the species which was being
rapidly exterminated left the sunshine and the enemies
which swarmed in it, and sought the ill-lighted recesses
of the bank. Of course, it is really a problem whether
these bees actually left the sunshine and crept away, or
whether the parasites had merely exterminated all those
which persistently remained in the sunny areas. With
nations of bees—as with nations of men—there is not
place in the sun for all. The numerous and the mighty
occupy the favored places, and any others that wish to
live may occupy the remaining corners of the earth until
such a time as the powerful wish to usurp the places in
the shade also. Usually the only hope of the ultimate
survival of such oppressed species depends upon their
powers of adaptation. Thus run the histories of the sur-
246 Trans. Acad. Sci. of St. Louis
vival of the fittest, the one proving its fitness by re-
maining in the sunshine thereby accelerating its devel-
opment, but curtailing its life, and the other proving its
strength by continuing its existence in spite of tre-
mendous opposition. One man’s meat is another man’s
poison; one bee’s sunshine is another bee’s death.
We eagerly await further developments in this com-
petition, but at present we cannot forgo the temptation
to speculate upon it. Will the white-banded bees, thus
hard pressed, be finally exterminated? If so, what will
be the effect upon Anthophora in return? Perhaps by
that time they will have become so strong that, like the
oak and the ivy, they will be able to carry on prosper-
ously the work of the mining-bees and also maintain the
various parasites without seriously feeling the strain.
But again, if this happens, what will become of those
later parasites which depended upon the white-banded
bees, etc., those which, in fact, have killed the goose that
laid the golden eggs? They will be compelled either to
change their mode of living in some way, to migrate and
perhaps perish in the attempt, or to quit living. If, how-
ever, the parasitic Diptera adhere to their tendency to
confine themselves to sunny areas, and if at the same
time the white-banded bees can adapt themselves to their
dim corners where they have taken refuge, there is a pos
sibility that the white-banded bees may again increase
and at some time regain prestige. Of course, these last
Sentences are merely speculative, and yet it is by pre
cisely such cat and mouse methods as these that nature
lets her many species run on in constant and deadly
competition.
The Ecology of a Sheltered Clay Bank 247
(c) Relation to cold, cloudiness, darkness.
Up with the early morning sun, at 6:30 in fine weather
the mining-bees of all three species, the carpenter-bees
and the Monobia mud wasps were all heartily at work.
They began early and worked diligently while the sun
shone and then relaxed soon after midday as soon as
the bank was in the full shadow. I have said elsewhere
that these insects know no calendar but the thermome-
ter; again we might well say they know no clock but
the sunlight. I hope some day to find out just what is
these bees’ program of daily work when their nests are
in the normal light, i. e. not shaded by artificial struc-
tures during any part of the normal hours of sunlight.
While the action described above was their usual pro-
gram, they sometimes made exceptions to this habit.
These digressions I have been unable satisfactorily to
explain. Often during the early part of June the An-
thophora bees were seen at work at 5:30 a. m., and con-
tinued busy until sunset. Despite the fact that May 28
was cloudy, some 20 mothers were actively engaged in
carrying in orange-colored pollen. On the other hand,
When on June 16 the temperature dropped twenty de-
grees, their activities were greatly lessened, and they
did not venture out to resume their work until 9 o’clock
on the 17th. Also after cold nights or rainy mornings
and toward the end of the season, even when the sun
did shine, often they did not come out early, but waited
until the bank was thoroughly warmed up. Sometimes,
when a run of cold days came in early autumn (in this
detail I am speaking of the white-banded bees), they did
not get out to work for two or three days; suddenly they
Seemed to be seized by the impulse to work, and then
248 Trans. Acad. Sci. of St. Louis
they would come out even on cold and gloomy days and
work hard, striving to complete their task almost as if
they felt in some way that time and life were ebbing
fast.
It will perhaps lead to a clearer understanding if we
pass from these generalizations to the record of what
actually happened after the rainy afternoon of August
29 and other cloudy and cold days. I merely give ex-
cerpts from the original notes.
‘August 30. The rain fell until 2 p. m. yesterday,
after that hour no insects were abroad. The night was
rather cold and today it was 10 a. m. before the first
insect appeared about the bank. A black cricket poked
his head out of one of the bee-holes where he had spent
the night. Soon several wasps, Tachysphex terminatus,
were scraping away the soil about their nests. A few
specimens each of Ancistrocerus fulvipes, Monobia qua-
dridens and Xylocopa virginica, and one each of Try-
poxylon clavatum, Chlorion auripes, and three white-
banded bees, Entechnia taurea were at work at their va-
rious duties of nidification: these were all observed ab
11:30 a. m. and all seemed unusually industrious, as if
to make up for the time lost in yesterday’s rain. As it
neared noon, the white-banded bees appeared in greater
abundance, and the parasitic bee-fly, Argyromoeba tig-
rina, made its appearance. That the sunshine was a fac-
tor in bringing out the bees is evidenced by the fact that
only from the nests that had been warmed by the di-
rect sunlight had the bees crept out to work. The
mothers from the nests in sunny sites were the first to
be active in bringing loads of pollen, while those nests
not so favored showed no external signs of life. I have
previously stated that this species of bee, Entechnia
The Ecology of a Sheltered Clay Bank 249
taurea, will burrow without regard to the light, that it
chooses dark areas as well as well-lighted ones without
any apparent detriment to the work, but here it is ap-
parent how, especially after a cold night, the sun indi-
rectly influenced the amount of work done by warming
up the nest and routing the sleepers. After all we must
admit, that despite the fact that these bees often built
their nests back in the shadow of the porch, they seldom
did work when the sun had passed beyond the zenith,
leaving the bank more shaded. It is interesting to note
that in the afternoon, after the deeper shadows fell upon
the bank at about 2 p. m. the activities of several spe-
cies were at their minimum. An analogous condition
existed on dark or cloudy days. The insects behaved
much as the population of a normal site would at twi-
light. As the shadows lengthened, some spiders would
become active, thus adding the suggestion of night-
prowlers to the twilight setting.
“‘September 4. While I have elsewhere recorded that
as the sun warms up the bank, more and more of the
Entechina bees come out of their holes and begin work,
it is now my pleasant duty to record that this rule is
not absolute, for today the behavior is reversed. Yes-
terday they were working intensely at their digging;
now it is dull and dreary after the heavy rain in the
night, yet the bees have been busily at work carrying in
pollen. In so far as I can see, this reversal of behavior
can be due only to the fact that much rainy weather
lately has caused a set-back in their work, and they be-
&in to ‘realize’ that if ever they are going to get their
domestic duties done before Gabriel sounds his trumpet,
they will have to be up and doing despite weather con-
ditions.
i
250 Trans. Acad. Sci. of St. Louis
‘‘September 12. The last two or three nights have
been very cold and today at 9:45 the only life about the
bank is a lone Tachysphex terminatus, an unfortunate
caterpillar in the jaws of an ant-lion, and about ten gray
lizards. None of the white banded females are in evi-
dence, but in the garden back of the house, eight males
were this morning found huddled in the cups of as many
wild bindweed blossoms where they had spent the night.
The flowers had not closed, evidently because of the low
temperature, and these male bees found lodging there
all night, and even at ten o’clock in the forenoon, they
were in a torpid and lethargic condition, heavily laden
with dew. This discovery so late in the season also
shows that they are not shorter-lived than the females
as we have found for other wasps and bees. By noon
the bank has warmed up perceptibly, and a few of the
sun-loving insects are out; one Entechnia taurea hovers
over her nest; one cuckoo-bee, Chrysis coeruleus lurks
suspiciously about. At two o’clock the first carpenter-
bee, Xylocopa virginica, and an Ancistrocerus fulvipes
are diligently caring for their nests. It is 3 p. m. be-
fore the first carpenter mud-wasp, Monobia quadridens,
becomes active about the wood tunnels overhead, and
even the Chalcid parasites, Monodontomerus sp. lin
ger to this late date. These meager numbers of each
species are no doubt indicative of the entire population
at this time of the year. It is interesting, too, to note
that despite the fact that most of the activities have
heretofore ceased somewhere about two in the after-
noon, we now find these remaining mothers working far
into the late hours of the afternoon. The day is too
short for them; they have their tasks and are intent upoD
finishing them, and in such stress are not guided abso-
The Ecology of a Sheltered Clay Bank 251
lutely and blindly by instinct, yet they do modify their
behavior according to their needs. Whether the sun-
shine is likewise warming the lizards, or whether the
appearance of possible food excites them, I do not know,
but at any event these vertebrates, heretofore sluggish,
become animated and active at the same time as the re-
mainder of the inhabitants.
“‘September 23. Only a few stragglers representing
some of the more important groups of the inhabitants,
remain at this late date, and they are indeed few in
number. It is evident that the cold of autumn is a fac
tor in reducing their numbers, but we should like to
determine precisely to what extent this is the deter-
mining factor, or in how far the weather conditions
directly influence their longevity. If the optimum con-
ditions for their existence prevailed for several months
longer, it might retard but would not abolish their mor-
tality. In other words, to what extent is longevity in-
herent in the organism, and how far is death due to
some external calamity?
“October 3. Today the most of the few individuals to
be seen certainly give one the impression that they have
outlived their allotted time—‘yet is their strength labor—
and sorrow.’ Three survivors of EZ. taurea linger help-
lessly about, as if they had been overlooked by death
and did not know how to conduct themselves in the land
of the living, where they no longer belong. Indeed no
activity is now possible for them, because the flowers
from which they have gathered pollen are long since
withered. Two Ancistrocerus fulvipes mothers are
feebly finishing the closure of their nests. Two or three
Monobia mud-wasps are seen on the rafters above the
bank, stunned and half dead with the cold. The only
252. Trans. Acad. Sci. of St. Louis
inhabitant showing any semblance to normal activity is
the grass-carrying wasp, Chlorion auripes. Four moth-
ers have been busy all this dull, gray day carrying in
blades of grass to plug up their nests in the old bur-
rows of the carpenter-bees. All the creatures of the unit
are now practically ready for their long winter sleep.
Soon the bank, frozen or snow-covered, will look utterly
barren and lifeless, yet within that mass of earth is life
—potential life that in the warm days of spring will
burst forth with renewed vigor, ready to battle again in
sharp competition for the possession of a small portion
of the earth which they may call home.”’
With the approach of cold weather the carpenter-bees
crowd into their tunnels and quietly await death. This
I suspected one spring when I opened a nest in a piece
of wood which contained six adults, all dead. During
eold days they often remain in the cells, crowded close
together, and come out again when the day grows
warmer. I have seen them come out for a little while
as late as October 13. On another occasion in early
September, I watched in vain for two days, for the oc
cupants of a nest. Concluding at length that they had
joined the heavenly choir, I proceeded to investigate
with an ax. At the first blow, a loud chorus of buzzing
greeted me, and then one by one twelve bees, each an-
grier than the last, flew out of the tunnel. This startling
experience made me wonder if it is not possible that part
if not all of them, hibernate in this way instead of dying
with the first cold of autumn. It seems to me that in
mild winters or in the southern states they might easily
hibernate thus.
There is some evidence that the clearness of the light
may be a factor in the homing behavior of the mining
The Ecology of a Sheltered Clay Bank 253
bees. When they returned from the field on their reg-
ular day trips, each tumbled precipitously into its own
hole without exploring the region or seldom trying out
any of the burrows before finding its own. (Figs. 1 and
4 show how similar and how close together the burrows
were). But one evening between 7 and 7:30, when it
was almost dark and bees were returning home, many
hesitated and hovered about before the colony and even
entered several holes before finding their own. Two
factors may enter into this inability to find their hole,
viz., the condition of weariness and the failing light.
This seems to us a logical foregone conclusion, and it
would be quite superfluous to mention it except for the
fact that a certain school of investigators argue for a
sixth sense to serve these Hymenoptera in their homing
flight or the ability to return home as if by magic.
(d) Relation to rain.
All of the mining bees carried water for their excavat-
ing. Rain helped them substantially by filling the wagon-
ruts in the road and giving them a supply of water
nearby. When these puddles dried up, they were obliged
to go a long way for water, and in some cases they did
without water or stretched a gulletful over so much
earth in the burrow that the result was not plastie mud
that could be shaped into turrets, but crumbly, slightly
moist pellets that could only be kicked out of the hole.
The bees did not leave their burrows during a rainy
spell, and I have never yet seen a mining-bee return to
the nest during a storm. I did, however, watch five
Monobia mud-wasps come home during a heavy down-
pour. They came in at intervals, all dripping wet
and slow of flight, and crept into their holes. The
254 Trans. Acad. Sci. of St. Louts
carpenter-bees nesting in the same place did not come
home during the storm. They saved themselves a drench-
ing, and after an hour or so, when the storm had ceased,
they came flying blithely in. We have recorded else-
where in these pages that the rain drops on the leaves
near by were eagerly lapped up by 4. abrupta and used
in their work of excavating, and I should not be sur-
prised if drops of dew in the early morning were gath-
ered for the same purpose.
While we find evidence of a certain amount of de-
struction of life by rainstorms, insects of a great variety
beaten down by the rain, and their nests in the earth
cut open or buried deeply by the erosion of the soil, yet
the rain is not without its benefits also. It has been
noticed that immediately after a rain following a long
dry period, there was a surprising increase in the pop-
ulation of burrowing Hymenoptera. For a time I east
this aside as mere coincidence, until the evidence seemed
too strong to be thus discarded. Why this sudden ap-
pearance after the rain, and what is the direct action of
the rain upon their emergence? There is no physiolog-
ical connection, I am sure, but merely a physical relation.
During a period of drought the earth had become ex-
tremely hard; hence the emergence of these young
ground-dwellers was rendered impossible. Perhaps they
could wait for a time, but surely not for long. We have
no way of knowing how many or what proportion of
them perished in their futile attempts to break through
the brick-like earth. The rain came and softened the
earth, and lo! they appeared as if by magic. No doubt
there was heavy elimination here, but in this case one
cannot call it the survival of the fittest; it was only the
survival of the lucky, and how do we know but the
The Ecology of a Sheltered Clay Bank 255
lucky ones in this case may have been the least fit to per-
petuate the race? In the activities of the turret-build-
ing bees also, we find wet or dry weather an important
factor in their nest building. In dry years, the turrets
are almost entirely omitted and in the case of A. abrupta,
when for the scarcity of water, much labor is put into
hard digging, the number of young is diminished.
(e) Relation of inhabitants to plant life.
At first thought, one sees but little relation between
the plant and the animal life of the bank. The vegeta-
tion about the base of the bank, scanty as it was, har-
bored numerous insects, such as small beetles, beetle
larvae and caterpillars, which sometimes dropped into
the ant lion pits. The grass-carrier wasp gathered grass
for her nest; the mining bee, the carpenter bee and the
Halictus went to the plants for nectar and pollen. The
caterpillar wasps A. fulvipes visited the plants for their
nectar and their caterpillars; the Trypoxylon and the
Tachysphex wasps went to the vegetation for their spl-
ders and grasshoppers respectively. As we have re-
lated before, the flowers of the wild morning glory served
so well as bachelor quarters for the males of the white-
banded bee that the females were obliged to fight for
even a chance to harvest the pollen from the flowers. The
bees came in from the surrounding vegetation bring-
ing pollen of various colors—yellow, orange, white, or
light green. One often found against the side of the
bank splashes of this pollen which had been lost in their
homeward flight.
For some time I suspected an intimate correlation
between the dates of the persimmon blossoms and the
emergence of the Anthophora bees. In the spring of
256 Trans. Acad. Scr. of St. Louis
1920, I watched to compare the dates. A large number
of Anthophora abrupta pupae which had been brought
into the laboratory early in April gave forth their young
in May;* the males emerged from May 10 to 25 and the
females only from May 22 to 25. On May 27, I visited
Wickes and found both sexes abundant about the clay
bank; this indicates that the time of emergence for the
controls in the laboratory was about the same as those
out-of-doors. Previously I had recorded the coincident
dying off of these bees with the dropping off of the per-
simmon blossoms, from which they gathered their pollen
and food. On May 27, I visited the permisson tree near
their colony, and found it laden with large buds, but
no insects were about the tree. The next morning, sev-
eral branches bore flowers; and lo! dozens of Antho-
phora bees were already present, joyously busy about
the new blossoms.
While this may at first seem to be only coincidence,
or to show that the bees had simply turned to any flowers
that they could find, we should not be hasty in dubbing it
mere chance until we have given due consideration to
the fact that the benefits of the association of blossoms
and bees may be mutual, and may be of as great impor-
tance to the one as to the other, or until we have more
data on these interrelations.
(f) Death by violence, and natural death.
To eat and to be eaten seems to be the law of the wild.
In the interrelation of life in the clay bank, we have re-
peatedly noticed how one insect falls prey to another.
We seldom think of natural death among the insects,
*Trans. Acad. Science St. Louis, 24 735, 1922.
The Ecology of a Sheltered Clay Bank 257
but this, too, occurs, and often to a much greater extent
than we usually suspect. During this study at the bank,
I have often picked up dead and enfeebled insects. Not
all the Chalcids fell prey to the spiders and ant lions;
I have removed hundreds from the crevices where they
had crept in to die. Many mining-bees were picked up
dead or limp at the end of their season and many more
were pulled out of their burrows. The cuckoo-bees were
picked up dead, too—and what enemy could wound a
cuckoo-bee in its thick armor? The blister-beetles, Hpi-
cauta marginata, dropped dead from the plants near the
base of the bank. Others found there whose death was
not accounted for, were Harpalus dichrous Dejean (E.
A. Schwarz), Tachysphex terminatus, Trypoaylon sp.,
Pseudagenia mellipes, Halictus pectinatus, Anthophora,
and Xylocopa virginica. Thousands of dead Chalcid
parasites were found in the burrows when they were
examined during the winter.
One often found newly dead female bees at the foot
of the bank. This was very puzzling, until one day I
actually saw where lay the trouble. A returning female
A. abrupta found that during her absence another bee
of the same species had appropriated her nest. A fight
ensued, in which the usurper was thrown bodily to the
ground and after a few twitches of the legs was dead.
Other fights of a less serious nature were often observed.
On one occasion I saw a female with part of her body
protruding from the burrow. I pulled her out with the
forceps, and found her tenaciously clinging with her
jaws to a second female that had evidently intruded in
the burrow. The next day also I saw the yellow, pollen-
laden legs protruding from a bee which seemed to be in
agony. I pulled it out with the forceps, and it, too, pug-
258 Trans. Acad. Sci. of St. Louis
naciously clung to another bee, which had evidently
taken advantage of the owner’s absence to usurp her
home.
(g) Relation to Soil conditions.
Not all of the soil that came out of the burrows went
into the making of the turrets. Much of it was kicked
out and fell on the ground below. This was neither dust
nor pellets, but characteristic minute bits or granules.
A very considerable heap of these granules had accu-
mulated in front of the center of the bank, where the
greatest activity occurred. This strip of drift meas-
ured 70 inches in length, 6 inches wide and from ¥% to
21% inches in depth. This of course gave the Tachysphes.
terminatus wasps an excellent medium in which to make
their nests, and afforded a material which made pit-
making for the ant-lions a pleasure.
It seems superfluous to mention the fact that, had the
nature of the soil been very different from what it was,
the colony of insects, especially the principal characters,
the mining-bees, would never have been there, since 4
firm, clayey soil of this nature is quite essential to mas-
onry of their kind; and too, if these pioneers had not
thrown out a large mass of granular soil to the bottom
of the bank, there would have been no ant lions, for these
never dig pits in firm soil.
(D)conc.upinG REMARKS,
After following the great things and the little things
in the lives of these tiny creatures, we may actually com-
pare the whole to a play upon the stage. The environ
ment corresponds to the stage; the dominant members
correspond to the leading characters, the secondary spe
The Ecology of a Sheltered Clay Bank 259
cies to the essential but subordinate characters. The
individual animals adjust themselves to one another,
and to the environment, as the personalities in the play
adjust themselves to one another and to the conditions.
‘‘In both groups some individuals are dominant, some
used and useful, some are tolerated, some pick up the
crumbs, still others are predatory or parasitic, and all
must be mutually adjusted to one another and to the
environment.’’
On this little stage of the clay bank we have seen en-
acted the tragedies and joys of life and love, birth and
death of many creatures; their tolerance of, or indiffer-
ence to one another, their craftiness in preying upon one
another and their alertness in escaping persecution, the
artisans at work, the visitors and the varying outcome
of their visits—all these things and many others per-
taining to the lives of a hundred insect species for a day,
a season, or a succession of years.
We have seen the pioneers blazing the trail, the rent-
ers following hard by, and the parasites ever on hand
keeping in check the otherwise rapidly increasing races ;
we have seen some that are transitory and some that
visit, prefer to remain, and may prove either a benefit
or a burden to the group.
And finally we have seen their behavior toward the
elements of nature, to sunlight and cold, to darkness
and to rain; all these little creatures would come and
go, act and react, conscious or unconscious of one an-
other and of the environment. All of the relations and
interrelations which have controlled them, and all of
the influence which they have wielded upon each other, to
bring this social unit to the degree of development or
balance in which we find it in the present years of the
260 Trans. Acad. Sci. of St. Louis
study, must have occurred within a short period, for
the existence of the clay bank covered less than a score
of years. How many bee kingdoms or families have
grown, lived and crumbled to oblivion here in that time,
we have no way of knowing. Under our very eyes we
have seen one grow to an unwieldy size at the expense
of another that has almost crumbled to decay.
It may seem naive for us to inquire into the mean-
ing of all the little commonplace phenomena which we
have observed here, and to search for many more not
yet observed. A casual reader might say ‘‘Of course
the organisms adjust themselves to each other and to
their environment; why all this fuss about it?’’ Per-
haps the same casual observer would have said to
Isaac Newton ‘‘Of course the apple will fall; why ask
questions about it??? And yet while we glibly call the
phenomena commonplace, we must admit that we really
know so little about the vital points in the lives of these
organisms in this one microcosm that if we were to at-
tempt to modify the balance of the population there we
should bungle the delicate adjustments so that our ex-
periments would not end anywhere near the point where
we aimed; and until we can thus direct the controlling
factors, we cannot boast an understanding of the sub-
ject.
‘‘Every reflective biologist must know that no living
being is self-sufficient, or would be what it is, or be at
all, if it were not part of the natural world, although no
truth is easier to lose sight of.’’ W. D. Brooks.
Sie ss
SeSnecease
F<
hee
ies
oe
Fig. 1.
Fig. 2.
Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
Puiate XIV
The work of An nthophora ey ag Not all hi bis excavated
soil went ie turret building, but much of i s kicked out,
and formed at the base (x) - medium for he habitel of ant-
lion larve. ,
The clay bank under the porch.
Trans. Acap. Sci. oF St. Lours, VoL. XXV PLATE XIV
264 Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
PLaTeE XV
Fig. 3. a ey ‘ewer of a ba bank. Note the absence of burrows
in contrast with figt
Fig, 4, ce EN. of the clay bank.
EUs. “VOU. AN
“AD. Sct. OF S
AC
TRANS.
266 Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
PLaTE XVI
Fig. 5. The mining-bee, Anthophora abrupta, (2 x natural size).
Fig. 6. The mining-bee, Anthophora raui, (2 x natural size).
Fig. 7. The white banded bee, Entechnia taurea, (2 x natural size).
Trans. Acap. Sci. oF St. Lours, Vor. XXV PiatTe XVI
268 Trans. Acad. Sci.‘of St. Louis
EXPLANATION OF PLATE
PLate XVII
2 the
Pig. 8: The “Ham and Bud” club. Note the clay bank under
south-half of the porch.
Trans. Acap. Sci, or St. Lours, Vou. XXV PuaTe XVII
9
-~
70
Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
Puate XVIII
Fig. 9. /The‘ turrets ia : AE eee E, Entechnia taurea; arrow,
Anthophora ra
Fig. 10. The u
was place
position.)
turned turrets of Anthophora raui. (A wad of cotton
d in the opening of each turret to show its up-turned
Trans, Acap:. Sci. or Str, Louts, Vou. AXV PLATE XVIII
272
Trans. Acad. Sci. of St. Lowis
EXPLANATION OF PLATE
Puate XIX
The Cells of Anthophora raui.
The pup of Anthophora raui,
A portion of the tunnel, and the orifice of the nest of Halictus
zephyrus.
Nymphs of Chortophaga_ viridfasciata.
Ant-lion pits in the dust at the foot of the bank.
Trans, Acap. Sci. or. Sr. Lours, Vou. XXV Pirate XIX
Trans. Acad. Sci. of St. Louis
EXPLANATION OF PLATE
PLaTE XX
A conspicuous inhabitant of the bank.
Gray lizards inhabited the bank.
Silver winged fly, Argyromoeba tigrina, (2 x natural size).
The shedding skins of Argyromoeba tigrina.
Trans. Acap. Sci. of Sr: Louis, Vout. XXV
PLATE XX
Trans. Acad. Sci. of St. Lowis
EXPLANATION OF PLATE
Pirate XXI
Fig. 20, 4 portion of the face of the clay bank, showing turrets
(E) Entechnia faurea, and (U) Anthophora abrupta. (Thi
shows the relative size of the turrets of the two species.
tu is horizontally made CE)' of penne taurea. (S)
Pig. 21; he
Pe upturned turrets of Anthophora ra
Fig. 22. Three years later; the same area as Fi ig 3; the nests of “st
abrupta are gradua ally spreading southward as the sunlig
shaded _ portions.
reaches the previously
Louris, VoL. XXV
1. OF Sr
TrANS. Acapb, Sc
FLORIDA FLOWERS AND INSECTS.
CHARLES RoBERTSON.
Except when Orlando is specified, the observations
recorded here were made at Inverness. Observations on
blooming seasons did not extend beyond April. Flowers
indicated as blooming the last of April probably bloomed
longer.
SIGNS, ABBREVIATIONS, ETC.
ab = abundant; ¢ = collecting pollen; esp = collecting
stray pollen; f = feeding on pollen; fq = frequent; fsp
= feeding on stray pollen; gn= gnawing; in cop = in
copula; lp = labial palps; Ma = non-social long-tongued
bee flower; Mas = social long-tongued bee flower; Mi =
non-social short-tongued bee flower; Mis = social short-
tongued bee flower; n= non-pollinating; 0 = ornitho-
philous; Pol = polytropic; R = red and all dark colors;
S = sphingophilous; se = sucking nectar and collecting
pollen; sf = sucking nectar and feeding on pollen; W =
white; Y = yellow, greenish to orange; ¢ = male; ? =
female; % — worker. The family ending ‘¢idae’’ is omit-
ted, as Halict for Halictidae. The months are Ja, F, Mr,
Ap, My, Jn, Jl, Au, S, O, N, D.
When no other indications are used after a name, the
insect is counted as sucking nectar legitimately and ef-
fecting pollination. Dates after ‘‘visitors observed’’ in-
dicate the period within which the observations were
made. Signs in parentheses, like ‘¢Halictidae (2, ¢)”’
mean that all of the following Halictidae were females
and all collecting pollen. Numbers in. parentheses, for
example (6,65), refer to the titles and pages in the
bibliography. In tables they are for species and visits or
(277)
278 Trans. Acad. Sct. of St. Louts
individuals. The plant names used here were taken from
Small, ‘‘ Flora of the Southern United States.’’
GENERAL SYNONYMICAL LIST.
The following is a list of all of the insects taken on
flowers, with authors’ names which are not repeated in
the special lists. Names which occur often enough to
justify it are abbreviated as follows: Bs. = Boisduval,
Cm. = Cramer, Cq. = Coquillett, Cr. = Cresson, Ed. =
Edwards, F. = Fabricius, L. = Linnaeus, Lp. = Lepele-
tier, Lw. = Loew, Mc. = Macquart, Mg. = Meigen, Rb.
= Robertson, Rh. — Rohwer, Ss. = Saussure, Sm. =
Smith, Tn.—Townsend, Wd.= Wiedemann, Wk. =
Walker, Wl. = Williston. Names in parentheses are the .
correct ones.
BIRDS.
Trochilidae: Trochilus colubris L.
LONG-TONGUED BEES (30).
Anthophoridae: Centris sp., Emphoropsis floridana
Sm.; Ap.:'Apis mellifera L.; Bomb.: Bombias scutellaris
Cr., B. separatus Cr., Siahais americanorum F., B. im-
patiens Har.: : Clergine- Ceratina dupla Say; Emphor.:
Melitoma tenes Say; Epeol.: Epeolus zonatus Sm.;
Eucer.: Florilegus condignus Cr., Melissodes sp., M. var-
iabilis Rb.; Megachil.: Ashmeadiella floridana Rb., type,
Coelioxqs 8. dentata Say, C. sayi Rb., Diceratosmia con-
junctoides Rb., type, Lithurgus ibboaie Sm., Megachile
floridana Rb. ie, M. generosa Cr., M. lanuginosa Sm.,
M. mendica Cr. , Sarogaster georgicus Cr.; Nomad: Cen-
trias rubicundus Oliv., Cephen fervidus Sm.; Stelid:
Dianthidium curvatum Sen D. (Anthidiellum) notatum
Latr., D. (A) perplexum fae Xylocop.: Xylocopa mi-
cans Los .. X. virginica Drury.
Florida Flowers and Insects 279
SHORT-TONGUED BEES (27).
Andrenidae: Opandrena scutellaris Rb., type; Collet:
Colletes sp., C. sp., C. americanus Cr., C. distinctus Sm.,
C. latitarsis Rb., C. thoracicus Sm.; Halic: ‘Agapostemon
splendens Lp., Augochlora fulgida Sm., A. sumptuosa
Sm., Chloralictus apopkensis Rb., type, C. ashmeadii
Rb., type, C. floridanus Rb., type, C. longiceps Rb., type,
C. nymphalis Sm., C. reticulatus Rb., type, C. tegularis
Rb., Evylaeus nelumbonis Rb., E. pectoralis Sm., Odon-
talictus capitosus Sm., Oxystoglossa sp., O. austrina Rb.,
type, O. matilda Rb., type, Panurg: Perdita obscurata
Cr.; Prosopid: Prosopis flammipes Rb., type, P. floridana
Rb., type, P. schwartzii Ckll.
OTHER HYMENOPTERA (100)
Bembicidae: Bembix spinolae Lp., Bicyrtes insidiatrix
Handlirsch, B. ventralis Say, Microbembex monodonta
Say; Cercer: Cerceris bicornuta Guerin, C. insolita Cr.,
C. austrina Fox, type, C. rufopicta Sm., C. verticalis Sm.;
Chalcid: Leucospis affinig Say, L. robertsoni Crawford,
type, L. slossonae Weld, Orasema sp. Spilochalcis flam-
meola Cr.; Chrysid: Chrysis intricata Brulle, Hedychrum
violaceum Brulle, Tetrachrysis venusta Cr.; Crabron:
Anacrabro robertsoni Rh., type, Hypocrabro decemma-
culatus Say, Solenius scaber Lp.; Ewmen: Eumenes fra-
ternus Say, E. smithii Ss. Leionotus apopkensis Rb.,
type, L. arvensis Ss., L. australis Rb., type, L. bicornis
Rb., type, L. bifurcus Rb., type, L. boscii Lp., L. floridanus
Rb., type, L. foraminatus Ss., L. fulvipes Ss., L. fundati
formis Rb., type, L. histrio Lp., L. histrionalis Rb., L.
megaera Lp., L. molestus Ss., L. oculeus Rb., type, L.
Saecularis Ss., L. turpis Ss., Monobia quadridens L.,
Odynerus erinnys Lp.; Larr.: Notogonidea argentata Bv.,
Tachysphex apicalis Fox, type, T. laevifrons Sm., Tachy-
tes aurulentus F., T. breviventris Cr., T. duplicatus Rh.,
280 Trans. Acad. Sci. of St. Louis
type, T. pepticus Say var. floridanus Rh., type, T. rob-
ertsoni Rh., type; Nysson: Nysson aequalis Patton;
Oxybel: Oxybelus floridanus Rb., type, O. fulvipes Rb.,
type; Pemphredon: Psen maculipes Fox, type; Philanth:
Philanthus carolinensis Banks, P. eurynome Fox, P. ven-
tilabris F.; Pompil: Allocyphonyx maurus Cr., Anoplius
illinoensis Rb., Aporinellus fasciatus Sm., Arachnoproc-
tonus ferrugineus Say, Ceropales robinsonii Cr., Episy-
ron biguttatus F., E. posterus Fox, type, Lophopompilus
philadelphicus Lp., Planiceps calcaratus Fox, type, P.
dubius Fox, type, P. minor Fox, type, Poecilopompilus
navus Cr., Pompiloides americanus Bv., P. argenteus Cr.,
P. marginatus Say, P. subviolaceus Cr., P. tropicus L.,
Psammochares tenebrosus Cr., Sericopompilus plutonis
Banks; Sphec.: Chalybion caeruleum L., Isodontia cin-
erea Fern., I. exornata Fern., Priononyx thomae F.,
Psammophila violaceipennis Lp., Sphex gracilis Lp., 8.
nigricans Dahlbom, S. pictipennis Walsh, S. procera
Klug, 8. vulgaris Cr.; Scoli: Campsomeris plumipes Dru.,
C. quadrinotata F., Discolia nobilitata F., Elis floridanus
Rh., type, E. propodialis Rh., type, E. robertsoni Rh.,
type; Tenthredin: Pseudosiobla robusta Kby.; Tpht:
Tiphia floridana Rb., type, T. vulgaris Rb.; Vesp: Polis-
tes americanus F., P. pallipes Lp., P. rubiginosus Lp,
Vespula maculata L.; Vipion: Cardiocheiles nigriceps
Viereck, Microbracon vernoniae Ash.
DIPTERA (104).
Agromyzidae: Milichia indecora Lw., M. robertsonii
Cq., type, Milichiella arcuata Lw.; Anthomy: Bithoraco-
chaeta leucoprocta Wd., Coenosia ovata Stein., C. sexno-
tata Mg., Homalomyia prostrata Rossi, Limnophora
narona Wk., Phorbia platura Mg.; Bibion: Dilophus
orbatus Say; Bombyli: Anthrax lateralis Say, ‘A. lucifer
F., Bombylius sp., Systoechus vugaris Lw., Systropus
Florida Flowers and Insects 281
macer Lw., Toxophora amphitea Wk., T. virgata O. S.;
Conop: Conops excisus Wd., Dalmannia vitiosa Cq.,
Physocephala sagittaria Say, Zodion nanellum Lw.;
Ephydr: Notiphila bicolor Cr., type, N. carinata Lw.,
Ochthera exculpta Lw.; Musc: Chrysomyia macellaria F.,
Lucilia caesar L., L. sericata Mg., L. sylvarum Mg,,
Musca domestica L., Synthesiomyia brasiliana B. & B.;
Ortal: Euxesta notata Wd., Tephronota humilis Lw.,
Oscin: Chlorops unicolor Lw., Siphonella cinerea Lw.;
Sapromyz: Pachycerina verticalis Lw.; Sarcophag: Heli-
cobia helicis Tn., Metoposarcophaga pachyprocta Parker,
Ravinia floridensis Ald., R. quadrisetosa Cq., Sarcophaga
assidua Wk., S. bullata Parker, S. incerta Wk., S. utilis
Ald., Sarcophagula occidua F.; Seps: Sepsis violacea
Mg.; Stratiomy; Nemotelus glaber Lw., Odontomyia
cincta Oliv., O. trivittata Say; Syrph: Allograpta obliqua
Say, Baccha clavata F., B. tarchetius Wk., Ceria signi-
fera Lw., Eristalis albiceps Me., E. dimidiatus Wd., HE.
transversus Wd., Helophilus divisus Lw., H. similis Mc.,
Mallota cimbiciformis Flln., Mesogramma boscii Me., M.
marginata Say, Microdon viridis Tn., type, Milesia vir-
giniensis Drury, Orthoneura nitida Wd., Paragus tibialis
F'illn., Psilota buceata Me., Spilomyia hamifera Lw.,
Syrphus americanus Wd., Tropidia albistylum Me., Volu-
cella fasciata Mc., V. sexpunctata Lw., V. vesiculosa F.,
Xylota pigra F.; Taban: Tabanus sparus Whitney;
Tachin: Archytas lateralis Mc., Atrophopalpus angusti-
cornis Tn., type, Atrophopoda singularis Tn., Chaeto-
glossa picticornis Tn., type, C. violae Tn., type, Cylin-
dromyia nana Tn., Ennyomma globosa Tn., type, Gonia
capitata De Geer, G. senilis Wl., Gymnoprosopa polita
Tn., type, Hypostena floridensis Tn., type, H. vander-
wulpi Tn., Masiphya brasiliana B. & B., Pachyophthal-
mus floridensis Tn., Phasioclista metallica Tn., Phasiop-
sis floridensis Tn., type, Phorocera edwardsii W1., Plagi-
Zaz Trans. Acad. Sct. of St. Louts
prospherysa parvipalpis Wulp, Polistomyia histrio Wk.,
Senotainia rubriventris Mc., S. trilineata Wulp, Siphocly-
tia robertsonii Tn., type, Siphona geniculata De Geer,
Siphophyto floridensis Tn., type, Siphosturmia rostrata
Cq., Spallanzania hesperidarum W1., Sturmia distincta
W4., Trichopoda pennipes F., Xanthomelana atripennis
Say; Trypet: Neaspilota vernoniae Lw., Urellia solaris
Lw.
LEPIDOPTERA (53).
Danaidae: Danaus archippus F., D. berenice Cm.; Hes-
peri: Achalarus lycidas 8. & A., Amblyscirtes vialis Ed.,
Ancyloxypha numitor F., Atalopedes cempestris Bs.,
Atrytonopsis loammi Whitney, Cocceius pylades Scudder,
Epargyreus tityrus F., Goniurus proteus L., Hylephila
phylaeus Drury, Lerodea eufala Ed., Megistias fusca G.
& R., Pamphila attalus Ed., Paratrytone aaroni Skinner,
Polites baracoa Lucas, P. brettus Bs., P. cernes B. & L.,
Prenes ocola Ed., Thanaos brizo B. & L., T. juvenalis F.,
T. martialis Scudder, T. terentius S. & B., Thorybes
daunus Cm.; Lycaen: ‘Atlides halesus Cm., Hemiargus
hanno Stoll, Strymon cecrops F., S. melinus Hbn.;
Nymphal: Dione vanillae L., Junonia coenia Hbn., Phy-
ciodes tharos Drury, Vanessa atalanta L., V. virginiensis
Drury ; Papilion: Papilio eresphontes Cm., P, glaucus L.,
P. marcellus Cm., P. philenor L., P. polydamas L., P-
polyxenes F., P. troilus L.; Pier: Catopsilia eubule L.,
Eurema delia Cm., E. euterpe Menetries, E. nicippe Cm.
Zerene caesonia Stoll; Rhiodin: Calephelis virginiensis
Gray; Satyr: Cissia sosybius F.; Sesi: Synanthedon
rubristigma Kellicott, S. geliformis Wk.; Arcti: Lerina
incarnata Bs., Utetheisa bella L.; Noctu: Psychomorpha
epimenis Drury; Syntom: Scepsis fulvicollis Hbn.
COLEOPTERA (15).
Buprestidae: Aemaeodera tubulus F.; Carab: Lebia
Florida Flowers and Insects 283
pumila De jean; Cerambyc: Typocerus zebratus F.; Chry-
somel: Donacia piscatrix Lac.; Cler: Trichodes apivo-
rus Germar; Dermest: Attagenus piceus Oliv., Orphilus
glabratus F.; Hlater: Cardophorus gagates Erichson;
Lampyr: Chauliognathus marginatus F., Polemius limba-
tus Lec.; Mordell: Mordella melaena Germar; Scarabae:
Euphoria sepulchralis F., Trichius (Trichiotinus) affinis
Gory, T. delta F., T. piger F.
HEMIPTERA (8).
Lygaeidae: Melanocoryphus facetus Say; Pentatom:
Euschistus crassus Dallas, E. servus Say; Phymat: Phy-
mata erosa L. var. guerini Lp. & S.; Pyrrhocer: Arhaphe
carolina H. Schaeffer, Largus longulus Stal; Reduvi:
Zelus cervicalis Stal, Z. bilobus Say.
When a genus contains only one species, the specific
name is omitted in the tables. Thus Apis = Apis melli-
fera &.
Insects were identified for me by J. M. ‘Aldrich, W. H.
Ashmead, Nathan Banks, D. W. Coquillett, J. C. Craw-
ford, E. T. Cresson, R. A. Cushman, W. J. Fox, G. H.
French, C. A. Frost, A. B. Gahan, C. A. Hart, Samuel
Henshaw, L. O. Howard, C. W. Johnson, Charles Liebeck,
J. R. Malloch, H. M. Parshley, R. R. Parker, Theo. Per-
gande, S. A. Rohwer, C. H. T. Townsend, P. R. Uhler,
and S. W. Williston.
Considerable difficulty has been found in getting the
insects identified. I have been compelled to meddle with
the business of the systematists and describe some bees
and Eumenidae, or go without names. This has put me
in the class of the jack-of-all-trades, giving the impres-
sion that I was collecting and describing bees from any-
where and everywhere, and I am offered specimens in
exchange for local ones.
284 Trans. Acad. Sct. of St. Louis
Sometimes insects sent by me for identification have
been retained and identified for the persons who retained
them, and it is often an accident that I find out the names.
A chalcid was identified for me as Leucospis distinguen-
dus and afterwards described as L. robertsont. A syrphid
identified as Microdon coarctatus was afterwards de-
scribed as M. viridis. The same specimen of Notiphila
was identified for me by three different dipterologists as
Dichaeta brevicauda, Notiphila carinata and N. bicolor.
One of these defended this practice because the correct
name had not been assigned. But that would not be fol-
lowed by any one who distinguished what he knew from
what he was guessing at.
The best way to get species identified is to describe
them yourself. One who will give you no aid in determin-
ing them will go 1000 miles to suppress them as synonyms.
Some author will describe species as if they differed
only in size and color and deposit the types in a distant
museum. You are not expected to read his descriptions
but must examine the types to see what he described.
His descriptions are ignored for many years. The spe-
cies are described under other names, which become as-
sociated with an extensive literature. Finally the types
are examined and familiar names are upset. The only
competent thing is for the systematists to clear up the old
names at first.
My experience is that the best time to get species iden-
tified is when an author is working on the group, when he
is competent to determine species or compare types. It
is hard to strike authors at that time. I have had proba
ble new species of Cerceridae lying in my collection while
three authors, one after the other, were working on them.
About the time you desire to send cercerids to one of
these, he is working on may-flies or spiders, like a flit-
Florida Flowers and Insects 285
ting schmetterling. I happened to strike Townsend when
he was working on Tachinidae and got many of them
identified, nine being types of new species.
Descriptions of bees in which the sexes are separated
as distinct species are worthless for identification.
Nomenclature is synonymical, not binomial. No one
has any authority to ignore priority*, but the historically
correct name often has no relation to the literature.
ACANTHACEAE.
CaLopHanes Ostonerroia, Ma., R.—The stems rise
1 dm. or less, and bear three or four flowers open
at a time. The corolla is 2 to 3 cm. long, expands
2m. The throat is 1 em. long and 7 to 8 mm. wide,
narrowing rather strongly to the tube which is 1 em.
long. The corolla is blue, but the throat below is marked
with purple. The flowers are nototribe and proteran-
drous. The throat is so wide that a bee which presses
its thorax against the anthers readily effects pollina-
tion. Butterflies can suck the nectar without much prob-
ability of effecting pollination. Mr. 15-Ap. 29, 17 spe-
cies, 100 individuals observed, Ap. 4-9.
Lone-roneurp Bess (4:14)—Anthophor.: Centris ¢ 1;
Bomb.: Bombus americanorum ? 1; Eucer. (2): Melis-
sodes sp. 1, M. variabilis 11. Swort-roncuep Bezxs
(3:5, ? )—Halict.: Agapostemon 1, Odontalictus 1, Oxy-
Stoglossa austrina 3, n. Orner Hymenoptera (1:1)—
“There are people who ignore history just as there are
people who legislate to make more than one-half of one
per cent of alcohol intoxicating instead of being appro-
priated as food, to limit a patient’s needs to one pint in
10 days, and to propagate the fake biology which 1 satis
ev:
olution. ea fd
286 Trans. Acad. Sci. of St. Lowis
Sphec.: Sphex procera 1. Drprera (1:12)—Bombyl.:
Systoechus 12. Lepmoprera (8:68, n.)—Hespert.: Lero-
dea 1, Pamphila 4, Polites baracoa 2, P. brettus 47; Nym-
phal.: Junonia 1; Papilion.: Papilio philenor 3, P. poly-
damas 3; Pier.: Eurema nicippe 7.
Ruetuia Hummus, Ma., R.—Mr. 12-Ap. 28, 3 visitors
observed, Mr. 14-26.
LeprpopTera (3)—Hesperi.: Pamphila; Papilion.: Pap-
ilio philenor, P. polydamas.
ANONACEAE.
Asmina Opovara, Mi, W.—Mr. 15-Ap. 26, 4 species
and 9 individuals observed, Ap. 8-14.
Conzorrera (3:8)—Cerambyc.: Typocerus 1; Scar-
abae.: Trichius affinis, 1, T. piger 6. Hemrprera (1:1)—
Pentatom.: Euschistus crassus 1.
APOCYNACEAE,
Amsonia Cruata, Ma., R.— The stem is terminated
by a conspicuous panicle of pale bluish flowers. The
corolla is tubular, with a border of 5 lobes. It meas
ures 14 mm. across. The tube is slender and measures
7 mm. in length. In the upper part, occupied by the
anthers and stigma, it is broader. The throat is densely
bearded, shielding the pollen from intruders.
The anthers surpass the stigma, so that self-pollina-
tion is not likely. The pollen is discharged in a mass-
The stigma is lateral and appears quite viscid. When es
insect inserts its tongue the viscid matter may readily
adhere to it and catch the pollen.
Long-tongued bees commonly show a mass of pollen 0?
their maxillary laminae, but butterflies never do. It is
doubtful whether butterflies ever earry the pollen. Mr. 8-
Florida Flowers and Insects 287
Ap. 26, visitors observed Mr, 18-Ap. 25. The following
5D species and 49 individuals carried pollen:
Lone-toneuep Bzzs (5:49)—Anthophor.: Centris ¢ 2,
2; Bomb.: Bombus americanorum °?, 6; Eucer. (82):
Florilegus 7, Melissodes sp. 18, M. variabilis 16.
The following 28 species and 112 individuals carried
no pollen:
Lone-roncuep Brzs (7:19)—Anthophor.: Centris 3;
Bomb.: Bombus americanorum 4; Eucer.: Florilegus
9, Melissodes sp. 1, M. variabilis 3; Megachil. ( ¢ ): Lith-
urgus 1, Megachile generosa 2. Oraer HyMENoPTERA
(3:5)—Sphec.: Isodontia cinerea 1; Scoli.: Camp-
someris quadrinotata 1, Discolia 3. Drerera (2:12)—
Bombyli.: Bombylius 1, Systoechus 11. Lepmoprera
(15:75)—Dana.: Danaus berenice 1; Hesperi.: Hylephila
3, Megistias 1, Pamphila 6, Polites baracoa 41, P. brettus
1, P. cernes 9, Thorybes 3; Nymphal.: Dione 1, Junonia 1;
Papilion.: Papilio cresphontes 1, P. marcellus 1, P. phile-
nor 4, P. troilus 1; Pier.: Catopsilia 1. Cotnoprera (1:1)
—Lampyr.: Chauliognathus 1.
ASCLEPIADACEAE,
Asciepias AmpLextcauuis, Mas, R.—Mr. 8-Ap. 26, 33
Species and 237 individuals observed, Mr. 19-Ap. 23.
The following 17 species and 66 individuals carried
Pollinia; h—on tarsal hairs, lp=on labial palps:
Lone-roncurp Bess (1:3)—Megachil.: Megachile gene-
rosa é 2, Ip, 3. SHort-roncuep Brees (2:2? )—Halict.:
Augochlora sumtuosa h, 1, Oxystoglossa austrina Ip, 1.
OrHer Hymenoprera (5:15)—Bembic.: Bembix h, 2,
Microbembex h, 1; Ewmen.: Leionotus foraminatus, on
claws, 1; Sphec.: Priononyx h, 10; Scoli.: Discolia h, 1.
(1:1)—Bombyli.: Systoechus, on pulvilli 1, Lept-
288 Trans. Acad. Sct. of St. Louis
poprera (8:45, h)—Hesperi.: Achalarus 1, Cocceius 1,
Pamphila 11, Polites baracoa 23, P. brettus 2, P. cernes 2,
Thorybes 4; Nymphal.: Junonia 1.
The following 172 individuals, belonging to 27 species,
carried no pollinia:
Lone-toncuep Bess (2:32)—Megachil.: Coelioxys 8-
dentata ¢, 1; Megachile generosa 31. SHonrtT-ToNGUED
Brzs (3:8? )—Halict.: Chloralictus floridanus 2, C. nym-
phalis 1, Oxystoglossa austrina 5. Orner HyMENOPTEBA
(7:26)—Bembic,: Bembix 3, Microbembex 2; Eumen.:
Leionotus arvensis 2; Larr.: Tachytes pepticus var. florl-
danus 2, type; Sphec.: Priononyx 15; Scoli.: Campsome-
ris plumipes 1, C. quadrinotata 1. Drerera (2:3)—
Conop.: Zodion 2; Taban.: Tabanus 1, Leprooprera (11:99)
—Dana,.: Danaus archippus 1, D. berenice 3; Hespert.:
Cocceius 2, Pamphila 9, Polites baracoa 11, P. cernes 1,
Thorybes 1; Nymphal.: Junonia 2; Papilion.: Papilio
marcellus 1, P. philenor 1; Pier.: Zerene 1. CoLEoPTERA
(1:2)—Lampyr.: Chauliognathus 2. Hemrprera (1:2)—
Reduvi.: Zelus cervicalis 2.
The Lepidoptera show the highest percentages “ gor
cies and individuals both with and without pollinia. OF
the species, 42.19%’ but of the individuals 31.2% show pol-
linia. Of Other Hymenoptera individuals, 36.5% show
pollinia. Only 11.1% of bee individuals show them. hs
most common bee, Megachile generosa, showed 34 in
duals, of which only 3 carried pollinia, and then only
on the labial palps.
Ascuepias Vertiomiata, Pol, W.—Mr. 4-17, one visitor
observed at Orlando, Mr. 4.
Orner Hymenoprera (1)—Tenthredin.: Pseudosiobla,
pollinia on legs, tongue and face.
Florida Flowers and Insects 289
AURANTIACEAE,
Cirrus Aurantium, Ma, W.—Visitors observed Mr. 31.
Lone-toneureD Bers (3)—Bomb.: Bombus americano-
rum ¢, sc, ab; B. impatiens ? 3; Xylocop.: Xylocopa
virginica 2. Orner Hymenoptera (3)—Larr.: Tachytes
aurulentus ; Scoli.: Campsomeris quadrinotata fq; Vesp.:
Vespula. Lepmoprera (3)—Nymphal.: Junonia; Pap-
thon.: Papilio philenor; Pier.: Catopsilia.
CACTACEAE,
Opuntia Austnina, Ma., Y. Mr. 30-Ap. 29, 15 species,
51 individuals observed, Ap. 7-25.
Lone-roneurp Bzzs (7:31)—Bomb.: Bombias scutella-
ris %, 1; Eucer.: Melissodes variabilis 3, se, 13;
Megachil.: ‘Ashmeadiella ¢, c, 1; Lithurgus ¢ 2, se, 13;
megachile generosa ¢?, 1, M. lanuginosa ¢, 1,
mendica ¢,1. SHort-roncuep Bzzs (2:4, 2, ¢)—Halict.:
Agapostemon 3, Chloralictus numphalis 1. Lzpmop-
Tera (4:5)—Hesperi.: Pamphila 1, Polites brettus 2;
Papilion.: Papilio philenor 1; Pier.: Eurema nicippe 1.
Couzoprera (2:11, f)—Mordell.: Mordella 1; Scarabae.:
Trichius piger 10, in cop.
CAPRIFOLIACEAE.
Lowicrra Sempsrvinens, Ma. (0) R.
Bros (1)—Trochil.: Trochilus ab.
Lepiwoprera (1)—Papilion.: Papilio cresphontes.
CISTACEAE,
Hewianroemum Carournianum, Ma., Y. F. 2-Ap. 29.
The stems rise a few inches and bear solitary, yel-
low flowers an inch or more in width. The petals are
obovate and are expanded horizontally. The numerous
290 | Trans. Acad. Sci. of St. Lows
stamens are also horizontal, their dehiscent anthers fac-
ing upwards. In this position there is no chance of the
pollen touching the stigma. The large stigma is sessile.
If a large bee loaded with pollen lands upon the flower
it must dust the stigma before touching the anthers.
Cross-pollination in this case is between different plants.
If the insect does not bear pollen, it may readily effect
self-pollination. The flower is visited exclusively for
pollen, the petals, as in most pollen-flowers, soon falling.
Females of small Halictidae, Chloralictus, Evylaeus and
Oxystoglossa, were observed collecting pollen.
COMPOSITAE.
Brruanpirera Susacauuis, Mas, Y.—The heads are
Single on the scapes and measure about one inch across.
There are 8 to 10 broad, yellow ray-flowers which
are pistillate. The lobes of the style are long, nearly
erect, with stigmatic edges. The disc florets have
nectar-bearing tubes 4 mm. deep. They are staminate,
the pollen being carried out on a long style brush, as in
Silphium. Ja. 10-Ap, 29, 14 visitors observed F. 27-Ap. 4:
Lone-tonevep Bzzs (2)—Megachil.: Coelioxys sayi °5
Megachile lanuginosa ¢. SHort-roncuep Buss (5, ?)—
Halict.: Augochlora sumptuosa, Chloralictus floridanus
sc, Evylaeus nelumbonis, E. pectoralis, Odontalictus s¢.
(2)—Bombyli.: Anthrax lucifer; Tachin.: Chae-
toglossa picticornis. Leproprera (5)— Hesperi.: Polites
baracoa, Thorybes; Nymphal.: Phyciodes; Pier.: Eurema
nicippe; Rhiodin. : Calephelis.
On ten days, Mr. 7-21, the following 300 individuals
were taken on flowers:
Lone-roneusp Bzzs (8)—Ap. 1, Epeol. 3; Megachil. 4.
SHort-ronevep Brrs (216)—Halict, 216. Drermra (9)—
Florida Flowers and Insects 291
Bombyli. 5, Syrph. 4. Lepmoprzra (67)—Arcti. 1, Hes-
peri. 43, Nymphal. 12, Papilion. 7, Pier. 4.
Odontalictus capitosus 2 , belonging to Halictidae, 8.3%
of the families represented, showed 65.3% of the indivi-
duals.
Carpuvus Sp., Mas, R.—One visitor observed at Orlando,
Mr. 21
Lone-roneuep Brzs (1)—Emphor.: Melitoma ¢ °.
_Lycopzsm1a ApHytia, Mas, R.—Mr. 21-Ap. 29, 5 spe-
cles and 15 individuals observed, Ap. 6-12.
Lone-ronevep Bzzs (1:1)—Megachil.: Lithurgus ¢, 1.
SHorT-toneurep Brzs (1:1, 2, ¢, n)—Halict.: Chloralictus
floridanus. Lepmoprera (2:12)—Hesperi.: Pamphila 6,
Polites brettus 6. Conzoprera (1:1, f, n)—Scarabae.:
Trichius affinis 1.
ERICACEAE,
Batopenpron Arsoreum, Ma, W.—The trees rise sev-
eral metres and are fairly white with flowers, which
are arranged in small racemes. The corolla is 4 mm.
wide at the thoat, so that the bodies of small insects
are admitted. The lobes are slightly reflexed, enabling
bees to cling to them more readily. The corolla is 7 mm.
long, its tube 5mm. The filaments have a pair of awns,
which when struck, cause a downpour of dry pollen.
Nectar is secreted by an epigynous disc. To reach it a
tongue must be thrust between the filaments. The latter
are hairy, which helps to protect the nectar and to cause
a pollen discharge.
The flowers are homogamous. Cross-pollination is se-
cured by the stigma being far in advance of the anthers.
292 Trans. Acad. Sci. of St. Louts
In their first visits bees may effect self-pollination. In
the absence of insects self-pollination may occur by the
pollen falling upon the slightly projecting rim of the
stigma. F’. 22-Ap. 24, 26 species and 68 individuals ob-
served, Mr. 16, 23, Ap. 5.
Lone-roneveD Bres (5:25)—Anthophor.: Empho-
ropsis ¢, sc, 7; Bomb.: Bombias separatus %, se, 1,
Bombus impatiens ? %,sc, 9; Xylocop.: Xylocopa micans
@, 1, X. virginica ¢ ¢°, 7. SHort-roncueD Berss
(3:7, 2 )—Halict. (sc): Agapostemon 5, Evylaeus pecto-
ralis 1; Prosopid.: Prosopis flammipes 1, type. OTHER
Hymenoptera (8:17)—Eumen.: Leionotus apopken-
sis 1, L. megaera 2, Monobia 5; Scoli.: Campsomeris
plumipes 2, C. quadrinotata 4; Vesp.: Polistes ameri-
eanus 1, P. pallipes 1, P. rubiginosus 1. Drprera
(2:3)—Musc.: Chrysomyia 1; Syrph.: Milesia 2. Leri-
DoPTERA (8:16)—Hesperi.: Cocceius 1, Epargyreus 1,
Goniurus 1; Thanaos juvenalis 1; Papilion.: Papilio cres-
phontes 2, P. marcellus 3, P. philenor 6, P. troilus 1.
Gayussacia Hrrrenza, Ma., W.—F. 17, 18, 9 species and
28 individuals were observed at Orlando.
Lone-ronevep Bess (1:1 2 )—Bomb.: Bombias separa-
tus 1. Swort-roncurp Brxs (1:5 ¢)—Halict.: Augo-
chlora fulgida 5. Oraer HyMEnoprera (7 :22)—Eumen.
(at holes, n): Eumenes smithii 1, Leionotus histrio 3,
Odynerus 7, perforating; Scoli.: Campsomeris plumipes
LG. ae ear ce! Q iat (at holes, n): Polistes ameri-
canus 4, P. pallipe
Preris Nirma, Ma., R.—The stems rise one OF
two metres and the wie of the branches are crowded
with pendulous flowers. The corolla is cylindraceous
Florida Flowers and Insects 293
with a gibbous base. It is from 7 to 9 mm. long and is
contracted in the throat to 1 or 2mm. The stamens and
stigma are included. The filaments are crumpled and at
the apex are two-awned, so that an entering tongue is
certain to touch them and shake out the loose pollen.
Blooms February 2-April 26, 1 visitor observed,
February 19.
Lepwoprera (1)—H espert.:; Thanaos juvenalis.
Viccintum Nitipum, Ma., W.—The corolla is white,
often with a reddish tinge, urceolate, 5 mm. long.
The mouth is so constructed that all except the smaller
insects are excluded, and for ordinary ones a tongue 5
mm. long is needed to obtain the nectar. The bee touches
the exserted stigma before the included anthers. The
anthers have scarious tips with terminal pores through
which the light pollen is sifted down upon the visitor.
Nectar is secreted by the thickened epigynous disc. Ja. 24-
Ap. 23, 18 visitors observed, F. 16-M. 7.
Lone-ronevep Brzs (2)—Anthophor.: Emphoropsis ¢
%, fq; Megachil.: Megachile lonuginosa ¢. SHorr-
TONGUED Brxs (1, 2 )—Halict.: Chloralictus longiceps.
Orner Hymenoprera (3)—Eumen.: Odynerus, perforat-
ing, n; Scoli.: Campsomeris plumipes fq, C. quadrinotata
fq. Lepworrmra (12)—Hesperi.: Cocceius, Polites bara-
coa, Thanaos juvenalis, T. martialis, T. terentius,
Thorybes; Nymphal.: Vanessa virginiensis; Papilion.:
Papilio marcellus, P. philenor, P. troilus; Pieri.: Eurema
nicippe, Zerene.
Xouisma Ferrucmvza, Ma., W.—The ends of the
branches are crowded with white flowers.
294 Trans. Acad. Sci. of St. Louis
The corolla is nearly globular, 3 or 4 mm. long by 4
mm. wide at the base and 2 mm. wide at the 5-toothed
throat. The 10 stamens and the style are included. The
anthers have terminal pores and hold their tips against
the style. A bee’s tongue disturbs the anthers and the
pollen is sifted out. The pollen does not fall upon the
stigma, though its edge may receive some.
The pendulous position favors bees, which easily hang
upon the flowers, but butterflies also effect cross-pollina-
tion. The mouth of the corolla is so narrow that only
bees with tongues about 4 mm. can drain the nectar.
F. 2-Ap. 24, 6 visitors observed, F. 2-Ap. 5.
Lonc-ronevep Bzes (3)—Anthophor.: Emphoropsis
3 2,8c, fq; Ap.: Apis; Bomb.: Bombus impatiens ? ¥-
Oruer Hymeyorrera (1)—Scoli.: Campsomeris quadri-
notata, Lxpipoprera (2)—Hesperi: Thanaos martialis;
Papilion.: Papilio marcellus.
ERIOCAULONACEAE.
Exrocauton sp., Mis, W—F. 7-Ap. 26, 9 species and 29
individuals observed, F. 29.
Lone-ronevep Bess (1)—Ceratin.: Ceratina ¢, 1.
SHort-roneurp Bugs (4:23)—Halict. (2): Chloralictus
ashmeadii 12, C. nymphalis 5, Evylaeus nelumbonis 1;
Prosopid.: Prosopis schwartzii ¢ 9,5. Drerera (3:4)—
Syrph.: Masogramma boscii 1, M. marginata 2, Tropidia
1. Lepwoprrera (1:1)—Nymphal.: Phyciodes 1.
EUPHORBIACEAE.
Cyiposcotus Strmutosus, Ma., W.—F. 8-Ap. 29, 2 visi-
tors observed, Mr. 19, Ap. 4.
Drerera (1)—Bombyli.: Anthrax lucifer. LepmoPTERA
(1)—Hesperi.: Thorbyes. Individuals observed Mr. 16,
17, 19, 21, were 15 Hesperiidae.
Florida Flowers and Insects 295
HYPERICACEAE.
Ascyrum TrrrapetaLum, Ma., Y.—The four yellow
petals are horizontal, the flower measuring an inch
across. The numerous stamens dehisce in _ succes-
sion. Since the stamens are turned more or less out-
ward, the anthers do not touch the stigmas. An insect
coming with pollen may readily effect pollination before
touching the anthers. Otherwise, it may effect self-pol-
lination. The flower seems to be visited only for pollen.
F, 2-Ap. 26, 4 visitors observed, F. 18-Mr. 23.
Lone-roneuzp Bes (1)—Megachil.: Megachile lanu-
ginosa ?, c, and n, cutting petals. SHort-ronauzp BEEs
(1)—Halict.: Chloralictus nymphalis °, ¢. Drerera (1)
—Syrph.: Mesogramma marginata f. Conzoprera (1)—
Buprest.: ‘Aemaeodera f.
LEGUMINOSAE.
Croratarta Rorunprrouia, Ma. Y. — The flowers
are separated on the racemes, so that only a few on
different stems are open at the same time. The insect
must land on each flower separately, and the flowers
must receive pollen from separate stems and commonly
from separate plants.
The flowers are yellow. The banner is nearly orbicu-
lar, measuring 8 mm. across and is erect or reflexed. Its
claw projects forward, and with the calyx serves to hide
the nectar so that a tongue 3 mm. long is needed to drain
it. At base the banner has cushions which serve the bee
as a head-rest when forcing down the keel. The wings
are short, of an orange red color, forming a path-finder,
and are folded close around the keel, so that the latter
dispenses entirely with its color. The base of the wings
lie upon the base of the keel and their tips so closely
envelope it that all are forced down together and, when
296 Trans. Acad. Sci. of St. Louts
released, return to their former position. The keel is
saccate at base, turns abruptly upward and ends in a
slender beak.
The stamens are monadelphous, a round opening in the
tube at base above allowing access to the nectar. When
the flower opens five oblong anthers have discharged
their pollen and five rounded anthers are still indehiscent.
In the bud the long anthers surpass the others. The five
stamens with long anthers rise to the tip of the keel,
shed their pollen, and then withdraw. The five stamens
with round anthers then lengthen, their indehiscent an-
thers serving to push the pollen up into the apex of the
keel. Then they shed their own pollen where it will be
easily carried out by the style-brush.
The style bends abruptly upward, and has a fringe of
hairs above and a circle of hairs about the tip. When
the keel is depressed, the style pushes out a ribbon of
pollen. If the keel be further depressed, the style itself
appears carrying out more pollen upon its brush. Ja. 21-
Ap. 29, 13 species, 90 individuals observed, F. 20-Ap. 20.
Lone-ronevzp Bzxzs (9:71)—Anthophor.: Centris ?, 1;
Megachil.: Ashmeadiella floridana ¢ 2, s¢, 13, type,
Diceratosmia ¢, 1, Megachile generosa ¢ 2, 9, M. lanu-
ginosa ¢ 2, sc, 20, M. mendica 2 , 7, Sarogaster ¢ %, 8¢
9; Stehd. (8 %, sc): Dianthidium notatum 4, D. per-
plexum 7. Dirrera (1:1, n)—Bombyli.: Systoechus 1.
Lepmwoprera (3:18, n)—Hesperi.: Polites baracoa 16,
Thanos brizo 1, Thorybes 1.
Eryrorina Herpacea, Ma. (0), R.—Several stems
rise 2 to 4 feet and bear scarlet flowers in racemes
1 to 2 feet long. The flowers are about 2 inches long:
The calyx tube is 10 mm. with short lobes. The wings
and keel are almost entirely included in the calyx tube,
Florida Flowers and Insects 297
serving merely as a protection to the stamens on the
lower side. The banner is folded longitudinally. At
base it enfolds the stamens, wings and keel and is itself
enfolded by the calyx tube. ‘Access to the nectar is on
each side of the free filament. The stamens are of
unequal length, so that the anthers form a pollen surface
12 mm. in length. They are enclosed in the infolded ban-
ner, but their tips are slightly exposed below. Visitors
must have a proboscis 14% inches or more to reach the
nectar conveniently. The stigma stands among the an-
thers and may easily receive their pollen, but the way
neglected flowers fall seems to show that self-fertiliza-
tion does not occur. It is evidently adapted to humming
birds, and dusts their throats with pollen. F. 26-Ap. 29.
Brrps (1)—Trochil.: Trochilus ab.
Lupinus Dirrusus, Ma., R.—Often the plants grow in
clusters a few feet across, which increases the conspicu-
ousness of the flowers. The racemes are erect and densely
flowered.
The flower is 15 mm. long. The banner has its blade
entirely exposed, since it has mainly an attractive func-
tion. At base it runs forward and is creased so as to
give a head-rest to bees depressing the keel. It is blue
with a longitudinal white spot, which forms a path-finder.
It is nearly orbicular, 12 mm. across.
The wings form a colored envelope of the keel. They
are coalescent at tip and move with the keel so as to keep
it concealed.
The keel is faleate, without color, and is bent abruptly
upward, ending in a sharp beak, as in Crotalaria. The
keel petals are coalescent above and below, leaving a
small opening at the tip.
298 Trans. Acad. Sct. of St. Louis
The absence of an opening in the stamen tube shows
that nectar is wanting, and that the flowers are adapted to
female nest-making bees, but they are visited by other
insects. In the bud the oblong anthers dehisce, filling the
tip of the keel with pollen. When the flower opens, the
round anthers are still closed. They no doubt aid in
forcing the pollen of the long anthers into the tip of the
keel and then dehisce.
The tip of the style is provided with a brush of hairs,
which, when the keel is depressed, forces the pollen out
of the tip of the keel. A long ribbon of pollen first ap-
pears and then the tip of the style. The style returns
to its former position when the keel is released, and then,
when the keel is forced down, again sweeps out the pol-
len. F. 7-Ap. 26, 29 species, 93 individuals observed,
Mr. 5-Ap. 5.
Lone-roneuzp Brzs (10:35)—Anthophor.: Emphorop-
sis 2, se, 1; Ap.: Apis sc, 2; Bomb.: Bombus american-
orum 2,1; Megachil.: Coelioxys octodentata ¢, 1, C. sayl
é ?,5, Megachile floridana 3,1, type, M. generosa ¢ %,
7, M. lanuginosa 3 9, s¢.,7 , M. mendica ¢ 2, sc., 8, Saro-
gaster 6,2. Sworr-roneuep Buss (2:9)—Halict.: Augo-
chlora sumptuosa 2°, ¢., 2, Chloralictus floridanus 2, fsp-,
7, type. Leproprera (17:49, n.)—Hesperi.: Achalarus 1,
Cocceius 2, Hylephila 1, Lerodea 6, Megistias 1, Pamphila
3, Polites baracoa 2, P. brettus 7, Thanaos juvenalis 1, T.
martialis 1, Thorybes 13; Lycaen.: Strymon melinus 1;
Nymphal.: Junonia 2, Vanessa virginiensis 1; Papilion.:
Papilio philenor 5, P. troilus 1; Pier.: Eurema nicippe 1.
T1um Inroysum, Ma., Y.—The stems are prostrate, the
short, dense-flowered racemes only turning up from the
ground. The flower is pale yellow and measures 11 mm.
The calyx tube measures 3 or 4 mm.
Florida Flowers and Insects 299
The banner runs forward for 5 mm., its claw, as well
as the claws of the other petals, being held by the calyx
tube. To reach the nectar a tongue 4 or 5 mm. long is
needed. The banner then bends upward, being con-
cave in front and strongly reflexed on the sides. As it
appears in front, it measures 7 mm. high and 4 mm. wide.
The folds render it quite rigid, so that it resists upward
pressure.
The wings are free, lying close to the keel, their tips
being bent to the right. This requires the bee to land
on the right side and, as in 7. obcordatum, pollen contact
is limited mainly to the left cheek. This slight disposi-
tion of the wings makes the flower pleurotribe. A
process in each wing fits in a pit in the keel, requiring
the parts to move together. On the base of each wing is
a hook-like process which clasps the stamen tube and pre-
vents the wings and keel from being thrust aside.
The keel petals are united. When the keel is de-
pressed, all of the stamens are exposed. When freed,
the keel again covers them.
The stigma receives pollen from the bee before the
anthers touch it. Self-pollination is hardly probable.
Access to the nectar is on each side of the free stamen.
F. 19-Ap. 16, 2 visitors observed, F. 25.
Lermorrera (2, n.)—Hesperi.: ‘Amblyscirtes, Polites
baracoa.
Trum Oscorpatum, Ma., R.—It grows flat on the
ground, and bears short racemes of pale-bluish flowers,
the tip of the keel being of a darker blue. The flower is
10 mm. long, the banner 5 mm. wide. The banner runs
forward for 6 mm., when it turns obliquely upward. At
base the banner clasps the other petals and is itself
300 Trans. Acad. Sct. of St. Lowis
clasped by the calyx, so that it resists the entrance of
small insects and limits the visitors to those having a
tongue 5 or 6 mm. long. On the base of the banner is a
large pale spot crossed by purple lines which run in-
wards, forming pathfinders. The keel petals are united.
A process in each wing fits in a depression in the keel,
so that keel and wings are depressed together.
Both wings turn to the right, which requires the bee to
land on the right. The bees taken on the flowers had
pollen on the left cheek, so that the flower is pleurotribe.
When the keel is forced down, the stigma and after it the
dehiscent anthers strike the bee, the stigma thus receiv-
ing pollen from other flowers. Self-pollination can
hardly occur if the flowers remains undisturbed. F. 16-
‘Ap. 23, 5 species and 19 individuals observed, F'.17-29.
Lone-toncugp Brzs (2:13, ¢ )—Megachil.: Diceratos-
mia 4, Megachile lanuginosa 9. Lxepmorrera (3:6, n.)—
Hesperi.: Amblyscirtes 3, Polites baracoa 1, Thorybes 2.
Viora AcutiForta, Ma., W.—The flower is 9 mm.
long, the abcordate banner 6 mm. wide. The banner is
pale bluish, marked in the middle with purplish streaks,
which, with the purple tip of the keel, form pathfinders.
Elsewhere the flower is white.
The banner has a broad base, enclosing the other
petals. It runs forward for 6 mm., thus limiting access
to the nectar. The wings lie ulonseido the keel, hiding it
from view, their tips far surpassing it. They are united
with the keel and move with it. The keel is short
(6 mm.), its petals united. Access to the nectar is on
each side of the free filament. When the keel is de-
pressed the anthers are exposed, but most of the pollen
is deposited on the style-brush. The style turns up and
Florida Flowers and Insects 301
is bearded near the tip. Mr. 9-Ap. 26, 4 visitors ob-
served, Mr. 11-Ap. 3.
Lone-toneurp Bers (1)—Anthophor.: Emphoropsis 2.
Lepworrera (3, n.)—Hesperi.: Polites baracoa; Nym-
phal.: Phyciodes; Pier.: Eurema euterpe.
LENTIBULARIACEAE,
Prinevicuna Exatior, Ma., R.—Mr. 16-Ap. 3, 2 visitors
observed at Orlando, Mr. 16.
Lone-ronevrp Brzs (2, 2, ¢., reversing)—Megachil.;
Megachile lanuginosa, M. mendica.
Pincurcuta Lurea, Ma., Y.—F. 19-Mr. 24, 2 visitors ob-
served, F. 19.
SHort-toncuep Brzs (2, ?)—Halict.: Agapostemon,
Odontalictus.
Pinevicuta Puma, Ma., W.—Each plant commonly
bears a single scape. Only one flower is open at a time,
so that cross-pollination between distinct plants is usual.
The corolla varies from white to bluish-purple. The
white form has the tube yellow within, 4 or 5 reddish
Streaks on the upper wall forming pathfinders. The pur-
plish form has the tube darker purple within and the
Same pathfinders. The corolla is 10 to 16 mm. long and
7 to 10 mm. wide. The tube measures from 5 to 7 mm.,
the spur 4 to 5 m., so that a tongue 9 to 12 mm. long is
needed to drain the spur. The tube of the corolla is only
2 or 3 mm. wide.
The stigma is just above the mouth of the spur, where
the proboscis of the visitor is certain to strike it. The
lower wall of the tube is folded inward longitudinally
and is covered with glandular hairs, requiring the in-
302 Trans. Acad. Sci. of St. Louis
sect’s tongue to pass nearer the upper wall and so in
contact with the stigma. The stigma is a valvular flap,
turned backward over the two anthers. When a but-
terfly withdraws its tongue the stigma is turned forward
and the tongue comes in contact with the anthers.
The flower is nearly erect and its border almost hori-
zontal. It remains open all day. It may also be visited
by night-flying lepidoptera. J. 24-Ap. 5, 10 visitors ob-
served, F. 18-23.
SHort-roncurp Brzs (1)—Halict.: Evylaeus pectoralis
@. Lepmwoprrera (9)—Hesperi.: Amblyscirtes, Lerodea,
Megistias, Polites baracoa, Thanaos juvenalis ; Nymphal.:
Phyciodes; Papilion.: Papilio philenor; Pier.: Eurema
nicippe, Zerene.
Urrrcunara Inriara, Ma., Y.—The scape is supported
on the water by a whorl of leaves and bears sev-
eral flowers. The corolla is 18 mm. long and 9 to 12 mm.
wide. The upper lip is 11 mm. long and 9 mm. wide.
In its basal concavity is situated the stigma and two
stamens. The lower lip has a palate which rises 5 mm.
above its base and shuts against the upper lip, conceal-
ing stamens and pistil and closing the spur. The flower
is yellow, some reddish streaks on the palate forming
path-finders. Below, the lower lip is three-lobed and lies
in contact with the spur.
The spur is saceate, notched at the tip, has a large
mouth and is 8 mm. long. The valve-like stigma lies upo?
and conceals the anthers. Later it rises so as to expose
them. Nectar is secreted by the spur. The flowers Te
main open a few days. Cross-pollination results from
insect visits and self-pollination can hardly occur. Only
one or two flowers on the scape are open at a time. The
flower is evidently adapted to bees which know how to
Florida Flowers and Insects 803
depress the palate. Butterflies might steal nectar with-
out transferring pollen.
Plants were found rising from the bottom in 91% feet
of water in Tsala Apopka Lake. The flowers sometimes
open while they are still two or three feet below the sur-
face. After the flowers reach the surface the stems sep-
arate from the ground but remain anchored by surround-
ing plants. Ja. 7-Ap. 20, 5 visitors observed, F. 19.
SHORT-TONGUED Bezs (3, 2 )—Halict.: Chloralictus reti-
culatus, Evylaeus nelumbonis; Prosopid.: Prosopis
schwartzii. Lxepmoprera (2, n.)—Hesperi.: Thorybes;
Pier.: Catopsilia.
Urricutarra Susutata, Mi., Y.—The slender scape rises
3 or 4 inches and bears two or three flowers, only one of
which is open at a time; so that pollination is between dis-
tinct plants. Sometimes the plants form thin patches
quite attractive to insects.
The flower is 6 to 11 mm. long, the lower lip 5 to 8 mm.
wide. In a general way it resembles U. inflata. The
flower is entirely yellow. The spur measures 5 to 7 mm.
The lower lip forms a convenient landing place, and the
weight of the insect opens the spur.
When insect visits fail, self-pollination probably oc-
curs from the stigma curling up so as to touch the pollen.
Ja. 11-Ap. 4, 2 visits observed, F. 28, Mr. 2.
Lermoprera (2, n.) — Hesperi.: Polites baracoa,
Thanaos terentius.
MALVACEAE,
Sma Ruomerroua, Mi., W.—Ja. 11-Ap. 28, 1 visitor ob-
served, Mr. 22.
Lepmorrera (1)—Hesperi.: Polites baracoa.
304 Trans. Acad. Sci. of St. Louis
NYMPHAEACEAE.
Castauia Renirormis, Mi., W.—F. 17-Ap. 20, 3 visitors
observed at Orlando, Mr. 18. Ap. 2.
Suort-ronevep Bzzs (2)—Halict.: Agapostemon ?, ¢.;
Prosopid.: Prosopis schwartzii ¢ ?, f. Drprera (1)—
Ephydr.: Notiphila bicolor fq., type.
Little plants, growing in shallow water, with pedun-
cles 9-19 dm. long, and petioles 18-25 dm., probably be-
long to a different species. See Nymphaea odorata
(3,125). One visitor observed at Orlando, F’. 24.
Suort-roncuzep Bees (1)—Halict.: Evylaeus nelum-
bonis ?,¢.
Nympuaza ‘Apvena, Mi., Y. (3,123)—Ja. 11-Ap. 26, 6
visitors observed at Orlando, F. 18-Ap. 11.
SHort-roncuep Bees (3)—Halict. (2, se.) : Agaposte-
mon, Evylaeus nelumbonis ab.; Prosopid.: Prosopis
schwartzii . Drerzra (2)—Ephydr.: Notiphila carinata;
Syrph.: Helophilus divisus. Cotzoprera (1)—Chryso-
mel.: Donacia f., gn., in cop., ab.
OLEACEAE.
OsmaytHus Americana, Mis., W.—The ends of the
branches are full of flowers. The corolla with its 4
broad, revolute lobes measures 5 mm. across. The tube
is 2 or 3 mm. long and 2 mm. wide in the throat, so that
the nectar is quite accessible. F. 22-Mr. 31, 5 visitors
observed, F’. 22, 26.
Lone-roncuEp Brgs (2)—Anthophor.: Emphoropsis ¢ j
Xylocop.: Xylocopa virginica ¢. Orner HyMENoPTEBA
(1)—Vesp.: Polistes rubiginosus. LepmopTERA (2)—Ly-
caen.: Strymon cecrops; Papilion.: Papilio marcellus.
Florida Flowers and Insects 305
ORCHIDACEAE,
Isiprum Gracriz, Ma., W., Spiranthes gracilis in 7, 52—
F’. 17-Mr, 16, 1 visitor observed at Orlando, F. 17.
Lone-roncurp Bess (1)—Megachil.: Megachile lanu-
ginosa ¢, pollinia on maxillary laminae.
Lrmoporum Sp., Ma., R., Calopogon pulchellus in 1—F.
21-Mr. 16, 9 visitors observed at Orlando.
Lone-roneurp Bres (2, 2 )—Bomb.: Bombias separa-
tus n.; Megachil.: Megachile lanuginosa with pollinia.
SHort-roncurp Bzzs (6, ? )—Halict.: Augochlora fulgida
with pollinia, A. sumptuosa with pollinia, Chloralictus
ashmeadii n., Evylaeus nelumbonis n., Odontalictus n.,
Oxystoglossa sp., n. Orner Hymenoprers (1)—Eumen.:
Leionotus oculeus n., type.
POLYGALACEAE,
Potyeata Nana, Ma., Y.—The flowers grow in close
spikes. The early ones are sessile and their flowers ap-
pear to be habitually self-fertilized. The later spikes are
borne on stems two or three inches high. The flowers are
pale yellow, the tip of the keel more orange-yellow.
The keel bears a crest of 8 slender lobes. Pollen is de-
posited on a terminal tuft on the style. The stigma is
lateral. Nothing seems to prevent pollen from the
stamens being carried down to the stigma, though in-
Sects will no doubt effect cross-pollination. A probos-
cis 4 mm. long can easily exhaust the nectar. Ja. 26-
Ap. 24, 1 visitor observed, Mr. 2.
Lone-ronevep Bzzs (1)—Megachil.: Megachile lanu-
ginosa 3.
306 Trans. Acad. Sci. of St. Louis
Ponyeata Ponyeama, Ma., R.—Mr. 22-Ap. 29, 3 visitors
observed, Ap. 20.
Lone-roneueD Brrs (1)—Stelid.: Dianthidium per-
plexum ?.
SHort-ToneveD Begs (1)—Halict.: Oxystoglossa aus-
trina ?.
Lepipoprera (1)—Hespert.: Pamphila.
POLYGONACEAE,
THYsANELLA Fiwperata, Mis., W.—Ninety-six species,
5999 individuals observed, N. 11-21. The Lower
Hymenoptera are 48.9% of the species and 42.5% of the
individuals, while the bees are 17.7% of the species and
40.5% of the individuals. Odontalictus capitosus, 1% of
the species, showed 17.8% of the individuals.
Lone-roncurp Brzs (6:87)—Ap.: Apis sc., 78; Bomb.:
Bombus impatiens ¢, 3; Megachil.: Coelioxys sayi ¢, 1,
Megachile lanuginosa ¢ ?,3,M. mendica ¢,1; Xylocop.:
Xylocopa virginica ?, 1.
SHort-ronevep Brzs (11:156)—Collet. (2): Colletes
sp. se, 10, C. americanus 1; Halict.: Agaposte-
mon ¢, 12, Augochlora fulgida é, 1, A. sumptuosa $, 1,
Chloralictus floridanus 4 ¢, sc., 12, C. nymphalis 2, 5, C.
reticulatus ¢ 2,3, Evylaeus pectoralis 2, 1, Odontalictus
é 2,8¢., 107, in cop., Oxystoglossa austrina ¢, 3.
Orner Hymenorrmra (48:255)—Bembic.: Bembix 1,
Bicyrtes insidiatrix 3, B. ventralis 4; Cercer::
Cerceris bicornuta 1, C. insolita 4, C. verticalis 1;
Chalcid.: Leucospis affinis 3, L. slossonae 3; Crabron.:
Anacrabro robertsoni 1, type, Solenius 1; Eumen.:
Kumenes fraternus 5, E. smithii 3, Leionotus australis
1, L. bifureus 1, L. fulvipes 1, L. histrio 3, L. molestus 1,
L. oculeus 2, L. saecularis 8, L. turpis 1, Odynerus 21;
‘Florida Flowers and Insects 307
Larr.: Tachysphex apicalis 2, T. laevifrons 1, Tachytes
breviventris 1, T. duplicatus 1, type, T. robertsoni 1,
type; Nysson.: Nysson 3; Philanth.: Philanthus ecaro-
linensis 27, P. eurynome 1, P. ventilabris 8; Pompil.:
‘Allocyphonyx 48, Episyron posterus 1, Pompiloides
marginatus 1, P. tropicus 4, Sericopompilus 1; Sphec.:
Priononyx 11, Sphex gracilis 3, S. nigricans 1, 8. pic
tipennis 1, 8. procera 6, S. vulgaris 1; Scoli.: Camp-
someris plumipes 2, Elis floridanus 36, type, E. propo-
dealis 6, type, E. robertsoni 3, type; Vesp.: Polistes amer-
icanus 14, P. rubiginosus 1; Vigion.: Cardiocheiles 1.
Diprera (13:34)—Bombyli.: ‘Anthrax lateralis 1, A.
lucifer 1, Systropus 2; Conop.: Conops 1; Sarcophag.:
Helicobia 1; Syrph.: Baccha clavata 6, Orthoneura 1,
Volucella fasciata 6; Tachin.: Archytas 5, Gonia capi-
tata 2, Phasiopsis 1, Siphosturmia 2, Trichopoda 5.
Lerioprera (16:57)—Hesperi.: Goniurus 4, Hylephila
3, Lerodea 1, Polites baracoa 11, P. brettus 2; Lycaen.:
Hemiargus 21, Strymon cecrops 1, S. melinus 5; Nym-
phal.: Junonia 1, Phyciodes 1; Pier.: Catopsilia 1, Eure-
ma delia 2, E. nicippe 1, Zerene 1; Satyr.: Cissia 1;
Syntom.: Seepsis 1.
Cotgorrera (2:9)—Lampyr.: Chauliognathus 1; Scara-
bae.: Trichius delta 8.
Hemiprera (1:1)—Phymat.: Phymata 1, n.
RHAMNACEAE,
Czanoruus MicropHyiuvs, Pol., W.—Ja. 24-Ap. 14, 178
Species and 971 individuals observed Mr. 1-31.
The Lower Hymenoptera show 34% of the species and
42.2% of the individuals; the flies 33.7 and 29.5; the bees
14 and 17. The flowers are pretty decidedly sphecopoly-
tropic. The Diptera and Lepidoptera show a decline in
308 Trans. Acad. Sct. of St. Louts
percentage of individuals. The list yields types of 25
new species.
Lone-roncurep Brzs (9:32)—Epeol.: Epeolus ¢ 2, 3;
Megachil.: Coelioxys 8-dentata ¢, 2, C. sayi # 2, 9,
Megachile generosa 3, 3, M. lanuginosa ¢, 8, M. men-
dica ¢, 3; Nomad.: Cephen 2, 2; Stelid.: Dianthidium
eurvatum ¢,1, D. perplexum ¢, 1.
SHort-toncuep Bess (16:134)—Collet. (¢ ¢%): Col-
letes distinctus 6, C. thoracicus 40; Halict. (¢): Aga-
postemon 1, Augochlora fulgida 2, A. sumptuosa S8¢.,
21, Chloralictus apopkensis 1, C. ashmeadii 2, C. flori-
danus 27, C. longiceps 1, C. nymphalis se., 20, C. reti-
culatus 1, C. tegularis 1, Evylaeus pectoralis 1, Odon-
talictus 2, Oxystoglossa austrina 1, type; Prosopid.:
Prosopis floridana 7, type.
Orner Hymenorrera (60:410) — Cercer.: Cerceris
austrina 47, type, O. rufopinta 18, C. verticalis 8;
Chalcid.: Leucospis affinis 5, L. robertsoni 9, type, Ora-
sema 2, Spilochalcis 1; Chrysid.: Chrysis 2, Hedychrum
6, Tetrachrysis 6; Crabron.: Hypocrabro 5, Solenius 28;
Eumen.: Eumenes fraternus 8, E. smithii 7, Leionotus
apopkensis 3, type, L. australis 9, type, L. bicornis 9,
type, L. bifureus 7, type, L. boscii 9, L. foraminatus 7,
L. fulvipes 1, L. fundatiformis 1, type, L. histrio 9, L.
histrionalis 9, L. megaera 1, L. molestus 1, L. oculeus
33, L. saecularis 41, L. turpis 11, Monobia 2, Odynerus
6; Larr.: Tachysphex apicalis 2, type, T. laevifrons 6,
Tachytes aurulentus 4; Oxybel.: Oxybelus floridanus 2,
type; Pemphredon.: Psen maculipes 2, type; Philanth.:
Philanthus eurynome 22; Pompil.: Allocyphonyx 2;
Aporinellus 1, Arachnoproctonus 1, Ceropales 1, Episy-
ron biguttatus 9, E. posterus 5, type, Planiceps caleara-
tus 3, type, P. dubius 1, type, P. minor 1, type, Poecilo-
pompilus 2, Pompiloides argenteus 1, P. marginatus 4,
Florida Flowers and Insects 309
P. subviolaceus 5, Psammochares 1; Scoli.: Campsomeris
plumipes 8, C. quadrinotata 1; Sphec.: Isodontia exor-
nata 2, Psammophila 1, Sphex vulgaris 8; Tiphi.: Tiphia
floridana 4, type; Vesp.: Polistes americanus 2, P. ru-
biginosus 1; Vipion.: Microbracon 1.
Diptera (60:287)—Agromyz.: Milichia indecora 21,
M. robertsonii 2, type, Milichiella 4; Anthomy.: Bithor-
acochaeta 1; Bibion.: Dilophus 1; Bombyli.: Anthrax
lateralis 21, A. lucifer 1, Bombylius 17, Toxophora am-
phitea 11, T. virgata 2; Conop.: Conops 4, Dalmannia 1;
Ephydr.: Ochthera 1, pred.; Musc.: Chrysomyia 2, Syn-
thesiomyia 1; Sapromyz.: Pachycerina 2; Sarcophag.:
Helicobia 9, Metoposarcophaga 1, Ravinia floridensis 1,
R. quadrisetosa 15, Sarcophaga assidua 1, S. bullata 1,
S. incerta 1, 8. utilis 1; Syrph.: Allograpta 2, Baccha
tarchetius 1, Mesogramma boscii 1, M. marginata 5,
Orthoneura 2, Paragus 2, Spilomyia 1, Volucella fasciata
6, V. vesiculosa 1, Xylota 2; Taban.: Tabanus 4; Tachin.:
Archytas 3, Atrophopalpus angusticornis 2, type, Atro-
phopoda 1, Chaetoglossa picticornis 7, type, C. violae 5,
Cylindromyia 3, Ennyomma globosa 4, type, Gonia sen-
ilis 4, Gymnoprosopa polita 36, type, Hypostena flori-
densis 6, type, Masiphya 1, Pachyophthalmus floridensis
6, type, Phasoclista 1, Phorocera 1, Plagiprospherysa 6,
Polistomyia 1, Senotainia rubriventris 11, S. trilineata 2,
Siphoclytia robertsonii 6, type, Siphona 13, Siphophyto
floridensis 10, type, Spallanzania 2, Sturmia 1; Trypet.:
Neaspilota 2, Urellia 4.
Lerwoprera (21:28)—Arcti.: Utetheisa 1; Hesperi.:
Achalarus 1, Amblyscirtes 1, Cocceius 1, Lerodea 2,
Megistias 1, Pamphila 2, Polites baracoa 1, Thanaos
juvenalis 1, T. martialis 1, Thorybes 2; Lycaen.: Stry-
mon cecrops 1, S. melinus 1; Nymphal.: Junonia 2, Phy-
ciodes 1, Vanessa virginiensis 1; Papilion.: Papilio
310 Trans. Acad. Sci. of St. Louts
troilus 1; Per.: Eurema euterpe 1; Rhiodin.: Calephelis
4; Sest.: Synanthedon rubristigma 1, S. geliformis 1.
Coxteoptera (7:39)—Cerambyc.: Typocerus 18; Cler.:
Trichodes 1; Dermest.: Attagenus 12, Orphilus 4; Elater.:
Cardiophorus 1; Lampyr.: Chauliognathus 1; Scarabae.:
Euphoria 2.
Hemiprera (5:41)—Lygae.: Melanocoryphus 1; Penta-
tom.: Euschistus crassus 7; Pyrrhocor.: Arhaphe 2,
Largus 22, in cop.; Redwvi.: Zelus bilobus 9.
ROSACEAE,
Amyepauus Persica, Ma., R.—Sixteen visitors observed,
. 10.
Lone-roncurp Brzs (1)—Anthophor.: Emphoropsis ¢
?, fq. Suorr-roneuep Brzs (3, 2° )—Halict.: Chloralic
tus longiceps, C. nymphalis, C. tegularis fq. Oruer Hy-
MENOPTERA (4)—Humen.: Leionotus histrio, Odynerus
fq.; Scoli.: Campsomeris plumipes ab., C. quadrinotata.
Dirrera (4, n.)—Agromyz.: Milichiella; Ortal.: Huxesta;
Sarcophag.: Helicobia; Seps.: Sepsis. Leprnoprera (4)—
Dana.: Danaus archippus; Papilion.: Papilio philenor, P.
polyxenes; Pier.: Catopsilia.
_Lavrocerasvs Carouinsana, Mis., W.—First 1000 indi-
viduals taken on the flowers, F. 5-Mr. 4, follow:
Lone-roncurp Brzs (96)—Anthophor. 1, Ap. 9%.
SHort-toncump Bes (573)—Andren. 527, Collet. 21,
Halict. 25. Oren Hymenoprera (13)—Chalcid. 4, Ewmen.
7, Larr.1, Vesp,1. Dierera (288)—Acalyptratae 11, An-
thomy, 29, Bibion. 1, Conop. 1, Muse. 118, Sarcophag. 32,
Syrph. 74, Tachin, 22. Lepmorrera (30)—Rhopalocera
29, Heterocera 1.
Florida Flowers and Insects 311
Opandrena scutellaris ¢ 2, sc., showed 52.7% of the in-
dividuals and is probably more important than all of the
rest together.
Papus Sezotina, Mis., W.—F. 3-Ap. 2, 29 species, 102
individuals observed, Mr. 14, 15.
Lone-roneuEp Bess (1:15)—Ap.: Apis se. 15. Suorr-
TONGUED Buss (4:7, 2? )—Collet.: Colletes sp. 1; Halict.
(se.) ::Agapostemon 3, Chloralictus reticulatus 1, C. tegu-
laris 2. Orner Hymenoptera (3:14)—Sphec.: Chalybion
1; Scoli.: Campsomeris plumipes 4, C. quadrinotata 9.
Diptera (14:40)—Anthomy.: Limnophora 1; Bombyli.:
Bombylius 2; Conop.: Zodion 3; Musc.: Chrysomyia 20,
Lucilia caesar 1, Musca 1; Stratiomy.: Nemotelus 1, Odon-
tomyia cincta 1, O. trivittata 1; Syrph.: Allograpta 1,
Kristalis dimidiatus 1, Helophilus similis 1, Psilota 1,
Tropidia 5, in cop. Lepmoprera (6:23)—Hesperi.: Achal-
arus 2, Polites baracoa 10; Nymphal.: Junonia 2, Papi-
lion.: Papilio eresphontes 2, P. marcellus 2, P. philenor 5.
Cotzoprera (1:3)—Lampyr.: Chauliognathus 3.
Prunus Umpetiata, Mis., W.—Ja. 18-Mr. 16, 89 visi-
tors observed F, 8-14.
Lone-roncuzp Buss (5)—Anthophor.: Emphoropsis
é , ab.; Ap.: Apis. se., ab.; Bomb.: Bombias scutel-
laris @ ; Xylocop. (8 2): Xylocopa micans, X. virginica
s¢., fq.
SHort-ronauep Buss (11)—Andren.: Opandrena scutel-
laris 3 9 , SC, in cop., type; Halict. (¢): Agapostemon,
Chloralictus apopkensis se., type, C. ashmeadii se., type,
C. longiceps, type, C. nymphalis sc., C. reticulatus sc.,
type, C. tegularis sc., Evylaeus pectoralis sc., Oxysto-
312 Trans. Acad. Sct. of St. Lows
glossa matilda, type; Prosopid.: Prosopis schwartzii é ¢,
ab., in cop.
Orner Hymenoptera (13)—Chrysid.: Chrysis; Crab-
ron.: Solenius; Eumen.: Eumenes fraternus, EB. smithii
ab, Leionotus bifurcus, L. floridanus type, L. his-
trio ab., L. saecularis, Odynerus ab. ; Scoli.: Campsomeris
plumipes; Vesp.: Polistes americanus, P. pallipes, P.
rubiginosus.
Diptera (32) — Agromyz.: Milichiella; Anthomy.:
Coenosia ovata, ©. sexnotata, Limnophora, Phorbia;
Bombyli.: Anthrax lateralis; Conop.: Physocephala,
Zodion; Musc.: Chrysomyia; Oscin.: Siphonella; Sar-
cophag.: Helicobia, Ravinia quadrisetosa, Sarcophaga
assidua, Sarcophagula; Syrph.: Allograpta, Ceria, Eris-
talis albiceps, E. transversus, Helophilus similis, Meso-
gramma marginata ab., Orthoneura, Psilota fg., Syrphus,
Tropidia, Volucella fasciata, V. sexpunctata, Xylota fq.;
Tachin.: Archytas, Hypostena vanderwulpi, Plagipros-
pherysa, Siphona, Xanthomelana.
Lepmworrera (28)—Dana.: Danaus archippus ab.;
Hesperi.: Epargyreus, Goniurus, Lerodea, Paratrytone,
Polites baracoa, Thanaos juvenalis ab., T. martialis;
Lycaen.: Atlides, Strymon eecrops fq., S. melinus fq.3
Nymphal.: Dione, Junonia, Phyciodes fq., Vanessa ata-
lanta, V. virginiensis; Papilion.: Papilio cresphontes, P.
glaucus, P. marcellus ab., P. philenor ab., P. polydamus,
P. polyxenes, P. troilus fq.; Pier.: Catopsilia, Zerene;
Rhiodin.: Calephelis; Arcti.: Lerina; Noctu.: Pachy-
morpha
Rusvs Vintosus, Mi., W.—F. 13-Ap. 26, 6 visitors ob-
served, Mr. 13. The list is fragmentary.
Lerworrera (6) — Hesperi.: Pamphila, Thorybes;
Nymphal.: Junonia; Papilion.: Papilio cresphontes, P.
philenor ab., P. troilus.
Florida Flowers and Insects 313
RUBIACEAE.
GeLsEMiIum Sempervirens, Ma., Y. — The corolla
is 30 to 35 mm. long, its five-lobbed border ex-
panding to the same extent. The base is 25 mm. long.
From a narrow part below, which is 8 mm. long by 2 mm.
wide, it widens regularly to a throat which is 15 mm.
wide. The throat within is rich orange.
In the short-styled form the 4 lobed stigma rises 10
mm. The 5 filaments are attached to the corolla for 8
mm., their free ends exposing the anthers 6 mm. beyond
the mouth of the tube. The stamens form a central col-
umn, the anthers being extrorse. The bee touches the
stamens exteriorly and inserts its tongue between the
filaments. The stigma lobes project between the fila-
ments. The tube is narrowed by the filaments, the pistil
and the stigma. A tongue at least 10 mm. long is neces-
sary to drain the nectar. There is a tendency to sterno-
tribe zygomorphism. The flower is horizontal, the bor-
der vertical, the upper lobes somewhat reflexed. The
stamens are bent a little upward, and Emphoropsis lands
upon them and enters on the upper side of the flower.
In the long-styled form the stigma is less exserted than
the anthers in the short-styled form and the stamens are
longer than the pistil of that form. The tube is ob-
structed by the anthers. The style is declined toward
the lower side.
Self-pollination might be effected by insects in the
short-styled form. In the other form it can hardly occur.
D. 7-Ap. 3, 5 visitors observed, F. 3-26.
Long-tonauep Bers (1)—Anthophor.: Emphoropsis
é 2, ab. Lepmoprera (4)—Hesperi.: Goniurus; Papilon:
Papilio philenor; Pier.: Catopsilia, Eurema nicippe.
314 Trans. Acad. Sct. of St. Louis
Hovustonia Rorunpirouia, Ma., W.—The flowers often
appear as mats a foot or more in extent, and are quite
attractive to insects. The corolla is salver-form, with
four ovate white lobes, the tube yellowish. The tube is
6 mm. long and nectar is secreted at its base. |
In the short-styled form the stigma stands in the
throat, while the stamens rise 3 mm. above. The stamens
are separated (3 to 4 mm.), so as to apply pollen to the
heads or other parts of the visitors, where it will be read-
ily received by the stigma of the long-styled flower.
The long-styled form can only deposit its pollen on the
tongues of the visitors, where it will touch the stigma of
the short-styled form. The stigma rises 3 to 4 mm. above
the mouth, the anthers being included. The throat is
narrowed by the introrse anthers. Ja. 5 to Ap. 26, 28
visitors observed, Ja. 30-F 24.
SHorRT-ToNGUED Brzs (1)—Halict.: Agapostemon ¢.
Orner Hymenoptera (1)—Scoli.: Campsomeris plumipes.
Lepmortera (26)—Dana.: Danaus archippus; 7 espert.:
Goniurus, Hylephila, Lerodea, Polites baracoa, P. bret-
tus, Prenes, Thanaos juvenalis, T. martialis, T. terentius,
Thorybes; Lycaen.: Strymon cecrops, 8. melinus; Nym-
phal.: Dione, Junonia, Phyciodes, Vanessa virginiensis;
Papilion.: Papilio marcellus, P. philenor, P. troilus ;
Pier.: Catopsilia, Eurema delia, E. euterpe, E. nicippe,
Zerene ; Rhiodin.: Calephelis.
RUTACEAE.
Facara Cuava-Hercuuis, Mis., Y—Mr. 16-Ap. 16; first
1000 individuals taken on staminate flowers, Mr. 17-AP.
2, follow:
Lone-toneuep Brzs (171)—Anthophor. 3, Ap. 128,
Bomb. 3, Epeol. 1, Megachil. 28, Nomad. 3, Stelid. 3,
Florida Flowers and Insects 315
Xylocop. 2. SHort-ronavep Bees (683)—Andren. 32, Col-
let. 259, Halict. 192. Orner Hymenoprera (108)—Cercer.
2, Crabron, 8, Eumen. 37, Ichneumon. 1, Larr. 1, Nysson.
1, Philanth, 24, Pompil. 5, Scoli. 7, Sphec. 4, Tiphi. 10,
Vesp.8. Drprera (177)—Bombyli. 7, Conop. 37, Muse. 86,
Syrph. 15, Tachin. 32. Lepmorrera (51)—Rhopalocera
50, Heterocera 1. Cotroprera (6)—Lampyr. 6. Hemie-
TERA (4)—Pentatom. 4.
The Colletidae, about 2.7% of the families, showed
25.9% of the individuals. Colletes thoracicus had 256
individuals. The Colletidae, Halictidae and Apidae,
about 8.3% of the families, made up 57.9% of the indi-
viduals.
SCROPH ULARIACEAE.
InysantHEs GranprFrtora, Ma., R.—The flower is blue,
the lobes more or less white, especially at the tips. The
upper lip is small and cleft, the lower spreading, with
three large rounded lobes. On the lower wall of the
corolla the filaments of the two lower sterile stamens form
two ridges which are covered with yellow pollen-like
glands. The filaments have about the middle an awn-
like process.
The stigma lies under the upper lip. The two fertile
anthers are coherent and lie on the upper wall about 2
mm. below the stigma. In this position self-pollination is
fairly impossible. The insect’s tongue touches the stigma
before the anthers. The stigma is sensitive and closes
after contact.
When the tongue touches the anthers and is withdrawn,
it turns back the lower lobe of the stigma and closes it.
The tube is narrow, rendered more so by the sterile
filaments, perhaps entirely excluding short tongues and
requiring long tongues to incline to the opposite side
316 Trans. Acad. Sci. of St. Louis
where they will touch the anthers and stigmas. The tube
is about 6 mm.
The flower is nototribe, but insects may land on any
side and insert their tongues. F. 15-Mr. 23, 4 visitors
observed, F’, 21-Mr. 7.
SHort-toncuep Brzs (1)—Halict.: Chloralictus ash-
meadii 2, fq. Leprpoprera (3)—Hesperi.: Polites bara-
coa, Thanaos juvenalis; Nymphal.: Phyciodes.
Linarra Canapensis, Ma., R. (2, 228; 5, 585)— F. 17-Ap.
26, 31 visitors observed at Orlando and Inverness, F. 17-
Mr. 24.
Lone-Toneuzp Bzzs (6)—Ap.: Apis; Megachil.: Coeli-
oxys sayi 2, Megachile lanuginosa ¢ ?, M. mendica
2; Nomad.: Centrias 2 ; Stelid.: Dianthidium notatum
8. SHort-Toneuep oa (6, 2 )—Halict.: Agapostemon,
Augochlora fulgida, A. sumptuosa, Chloralictus longi-
ceps type, Evylaeus pectoralis, Odontalictus. OTHER
Hymenoptera (1)—Scoli.: Campsomeris quadrinotata.
Diptera (3)—Bombyli.: Toxophora amphitea; Syr ph.
(f., n.): Baccha clavata, Mesogramma marginata. Ler!
DOPTERA (15)— Hesperi.: Ancyloxypha, Atalopedes,
Lerodea, Polites brettus, P. cernes, Prenes; N: ymphal. :
Junonia,. Phyciodes, Vanessa virginiensis; Papilion.:
Papilio marcellus, P. philenor; Pier.: Catopsilia, Eure-
ma euterpe, Zerene; Arcti.: Utetheisa.
PENTSTEMON Hirsutus, Ma., W.—Mr. 30-Ap. i, 1 visi-
tor observed, Ap. 8.
Lone-Tonevep Bers (1)—Bomb.: Bombus america-
norum 2,
SOLANACEAE.
Puysauis Arenicona, Mi., Y.—Mr. 28-Apr. 29, 1 visitor
observed Ap. 6.
Florida Flowers and Insects S17
SHoRT-TONGUED Bzzs (1)—Collet.: Colletes latitaris 3.
Sotanum Nicrum, Ma., W.—F. 24-Ap. 29, 1 visitor
observed at Orlando, F. 24.
Lone-roncuep Brzs (1)—Bomb.: Bombus impatiens,
2,6,
UMBELLIFERAE,
Hyprocoryie Umpeixiata, Pol., Y. (4, 456, 459)—F. 27-
Ap. 26, 33 species and 65 individuals observed at Or-
lando, Mr. 15-20.
SHort-toneven Brzs (1:1)—Panurg.: Perdita ¢, 1.
Orner Hymenoptera (10:18)—Larr.: Notogonidea 4,
Tachysphex apicalis 1; Oxybel.: Oxybelus fulvipes 1,
type; Pompil.: Anoplius 1, Lophopompilus 1, Pompi-
loides americanus 4, P. argenteus 3, P. marginatus 1,
P. subviolaceus 1; Tiphi.: Tiphia vulgaris 1. Dr-
TERA (19:41) — Agromyz.: Milichiella 1; Anthomy.:
Bithoracochaeta 1, Homalomyia 1, Limnophora 4; Musce.:
Chrysomyia 2, Lucilia sericata 1, L. sylvarum 4, Musca
4; Ortal.: Tephronota 7; Oscin.: Chlorops 1; Sarcophag.:
Ravinia floridensis 1, Sarcophaga bullata 2; Syrph.:
Baccha tarchetius 1, Mesogramma boscii 5, M. marginata
2, Microdon viridis 1, type, Orthoneura 1; Tachin.:
Phasioclista 1, Senotainia trilineata 1. CoLEOPTERA
(3:5)—Carab.: Lebia 1; Lampyr.: Chauliognathus 3,
Polemius 1,
VERBENACEAE.
Lantana Camara, Mas., Y.—Three visitors observed at
Orlando, F. 24,
Lepworrera (3) — Beek: Thanaos juvenalis; Pa-
Pilion.: Papilio marcellus, P. philenor.
318 Trans. Acad. Sci. of St. Louis
Lantana Oporata, Mas., R.—Ja. 24-Ap. 11, 17 visitors
observed, F. 16-Mr. 19.
Drerera (1, f, n) —Syrph.: Mallota. Lepmoprera
(16)—Dana.: Danaus archippus; Hesperi.: Atrytonop-
sis, Cocceius, Lerodea, Megistias, Polites baracoa,
Thanaos juvenalis; Nymphal.: Junonia, Phyciodes, Van-
essa virginiensis; Papilion.: Papilio cresphontes, P. phil-
enor, P. troilus; Pier.: Eurema euterpe, E. nicippe,
Zerene.
VIOLACEAE.
Viota Lancrouata, Mi.. W.— The flowers are small.
The petals are white, the lower one streaked with purple,
lateral not bearded. The flower is not adapted to pollen-
collecting bees. The spur is short. From the tip of the
style a narrow passage leads to the base of the tube, a
distance of about 4mm. The stigma is beaked and only
surpasses the anthers a little, but enough to prevent self-
pollination.
Insects land on the upper petals and approach the nec-
tary from above. The flower is therefore sternotribe, the
place of pollen-contact being the under surface of the
proboscis. Ja. 19-Ap. 26, 16 visitors observed, Ja. 19-
F. 20.
SHort-ToncuEp Bzxzs (3, 2° )—Halict.: Chloralictus
apopkensis, C. ashmeadii, Odontalictus. DrprERa (3)—
Tachin.: Chaetoglossa picticornis, C. violae fa, ‘ype
Siphona fq. Leprmoprera (10)—Hesperi.: Amblyscirtes,
Cocceius, Goniurus, Megistias, Polites baracoa, Thanaos
juvenalis; Nymphal.: Phyciodes; Papilion.: P apilio
philenor; Pier.: Eurema delia, E. euterpe.
Florida Flowers and Insects 819
FLOWER CLASSES.
Ma.—Non-social long-tongued bee flowers (including 0)
are 07.1% of the flowers observed, receive 34.4 and 28.6%
of the total and the pollinating visits. The class re-
ceives 71% of the pollinating visits of long-tongued bees
and 38.2 of those of Lepidoptera. These are respectively
32.4 and 38.9% of the total visits to Ma.
Ma shows for colors R 40.6, Y 25.0 and W 34.3%, and
for visits to each color 49.1, 15.7 and 35.1%. Of the total
visits to R, Y and W, Ma receives 71.0, 44.3 and 18.9%.
Ma shows maxima of long-tongued bees, 70.5, short-
tongued bees 33.6, and Lepidoptera 48.9.
Of total non-pollinating visits, Ma shows 78.4%, of
those of Lepidoptera 88.3, flies 75.0, Lower Hymenoptera ~
71.4 and short-tongued bees 66.6%. Many insects get
pollen or nectar without effecting pollination.
Ma shows maxima of Trochil, Anthophor, Bomb, Eucer,
Megachil, Stelid, Halict, Bombyli, Hesperi, Nymphal,
Papilion, Pier, Scoli, long-tongued bees, total Lepidop-
tera.
Mi.—The non-social short-tongued bee flowers are
17.8% of the flowers observed and receive 6.2% of the
Visits. Lepidoptera are 57.4% of the visitors and short-
tongued bees 22.2. The Lepidoptera are not better
adapted but more common.
The colors are R 10, Y 30 and W 60%, and the visits
R 7.4, Y 16.6 and W 75.9. But the percentages of total
Visits to Mi, R, Y and W are 1.9, 8.4 and 7.4.
Mas.—The social long-tongued bee flowers are 10.7 %o
of the flowers and show 8.4% of the visits. Of the visits
to Mas, the Lepidoptera show 53.4%.
320 Trans. Acad. Sct. of St. Louis
Mis.—The social short-tongued bee flowers are 8.9% of
the flowers and show 26.4% of the visits. Of the visits
to this class, the lower Hymenoptera show a maximum
of 28.2%, the flies with 27.0 and the Lepidoptera with
23.1. Of the visits of the flies, lower Hymenoptera and
short-tongued bees, 39.9, 36.5 and 28.8 are to this class.
Mis shows maxima of Xylocop, Collet, Lycaen, Sphee,
Bembie, Crabron, Philanth, Syrph, Conop, Stratiomyidae.
Pol.—The polytropic flowers are 5.3% of the flowers
and receive 24.4% of the visits. Of the visits to this class,
the flies show a maximum of 37.2, the lower Hymenoptera
with 33.4. Of the visits of the flies and lower Hymenop-
tera 47.0 and 39.8% fall under Pol.
Cases referred to Pol on account of the species of vis-
itors may be shown to belong to Mis when the individuals
are counted. In Fagara clava-herculis the short-tongued
bees forming 9.3% of the families, were 48.5% of the
individuals.
Pol shows maxima of Eumen, Pompil, Oxybel, Tiphi,
non-aculeata; Tachin, other Calyptratae, total Muscoidea,
total flies, lower Hymenoptera, Coleoptera, Hemiptera.
Social and non-social flowers —While the social (Mas,
Mis, Pol) are only 25% of the flowers observed by me at
Inverness and Orlando, they receive 59.2% of the visits.
Of 437 flowers observed at Carlinville, Illinois (8, 158),
the social, 45.4% of the total, received 78.9% of 13,971
pollinating visits. The differences are 34.2 and 33.5.
Couors.
Red.—Red flowers are 32.1% of the flowers observed
and receive 23.8% of the visits. Of the visits to red, the
Lepidoptera made 49.2%. Red shows maxima of Eucer
and Megachilidae.
Florida Flowers and Insects Sal
Yellow.—Shows 25% of the flowers observed and 12.1%
of the visits. Of the visits to yellow, 23.8% are made by
flies and 23.8 by Lepidoptera.
White—Shows 42.8% of the flowers and 64% of the
visits. The lower Hymenoptera show 84.2, flies 76.7,
Coleoptera and Hemiptera 64.5, short-tongued bees 56.7,
Lepidoptera 54.9, long-tongued bees 41.1% under white,
all maxima. Of total visits to white, the Lepidoptera
show 27.9%, lower Hymenoptera 27.0, flies 23.2, short-
tongued bees 10.6, long-tongued 7.5.
General colors—As we pass from red through yellow
to white, the long-tongued bees show 38.2, 20.5, 41.1; Lepi-
doptera 36.1, 8.8, 54.9; short-tongued bees 25.9, 17.3, 56.7;
Coleoptera and Hemiptera 12.9, 22.5, 64.5; lower Hymen-
opetera 10.7, 5.0, 84.2; flies 8.3, 14.8, 76.7. Or long-
tongued bees and Lepidoptera 36.7, 11.9, 51.3; other in-
sects 13,3, 12.2, 74.4. The determining condition is that red
generally shows the deepest seated nectar and white the
shallowest. At Carlinville, Illinois, were observed 13,971
pollinating visits to 437 flowers, of which 30.2% were red,
30.6 yellow and 39.1 white. Long-tongued bees show 34.0,
31.4, 34.5; Lepidoptera 36.8, 23.7, 39.3; short-tongued bees
11.9, 35.8, 52.1; flies 7.7, 32.9, 59.3; lower Hymenoptera
7.5, 29.4, 62.9; Coleoptera and Hemiptera 6,3, 31.3, 62.2.
Long-tongued bees and Lepidoptera show 34.6, 29.2, 36.0;
other insects 8.9, 32.9, 58.0.
Species and individuals.—In the case of 18 species in
which the individual insects were taken as they came, bees
shows 24.9% of 477 pollinating visits and 37.4% of 2,679
individuals. In the case of 41 flower species observed at
Carlinville, Illinois, bees were 31.6% of 405 species, made
40.2% of 1,452 visits and were 57.4% of 7,391 individuals
(9: 70.)
322 Trans. Acad. Sct. of St. Louis
INSECTs.
Long-tongued bees.—In the case of 338 visitors and 867
visits, the long-tongued bees show 8.8% of the species
and 11.7% of the visits; 13% of the pollinating and 1.9%
of the non-pollinating.. The maximum is 70.5% under
Ma. For colors, they show 38.2% to red and 41.1 to white,
or 6.1 more and 1.7 less than the percentages of colored
flowers observed. The Megachil, Bomb and Anthophor,
33.3, 13.3 and 6.6% of the long-tongued bees, make 44.1,
14.7 and 11.7% of the long-tongued bee visits. These,
with Eucer and Stelid, have maxima under Ma. The
Xylocopidae show maxima under Mis.
Short-tongued bees——Being 7.9% of the species, these
show 11.9% of the visits; 12.4% of pollinating and 8.8
of non-pollinating visits. The maximum is 33.6 under
Ma. They show 25.9% to red and 56.7 to white, or 6.27
less and 13.9 more than the percentages of those colors.
All of their non-pollinating visits are to red flowers for
which they are too small.
The Halict, 59.2% of the short-tongued bees, make
85.5% of the short-tongued bee visits. They show maxima
under Ma. The Collet show maxima under Mis.
Lower Hymenoptera.—Being 29.5% of the visitors they
make only 20.5% of the visits; 21.4% of the pollinating
and 13.7 of the non-pollinating. A maximum of 39.8% is
under Pol. They show 10.7 under red and 84.2 under
white, or 21.4 less and 41.4 more than the percentages of
those colors. The Eumen., Pompil. and Scoli., 21, 19 and
6% of the lower Hymenoptera, make 27.5, 14.0 and 14.67
of the lower Hymenoptera visits. They show maxima as
follows: the Scoli under Ma; the Spec, Bembic, Crabron
and Philanth under Mis; the Eumen, Pompil, Oxybel,
Tiphi and non-aculeata under Pol.
Florida Flowers and Insects aac
Fles.—These are 30.7% of the visitors and make
19.3% of the visits; 20.3% of the pollinating and 11.7 of
the non-pollinating. The maximum is 47% under Pol.
They show 8,3 under red and 76.7 under white, 23.8 less
and 33.9 more than the percentages of those colors.
They show maxima as follows: the Bombyli under
Ma; the Syrph, Conop and Stratiomy under Mis; the
Muscoidea under Pol. They show in percentages of fly
species and visits: Bombyli 6.7, 10.7; Syrph 23.0, 26.7;
Tachin 27.8, 23.2; other Calyptratae 14.4, 16.0; total
Calyptratae 42.3, 39.1; total Muscoidea 61.7, 54.7.
Lepidoptera.—Are 15.6% of the species and make
82.5% of the visits; 29% of the pollinating and 58.8%
of the non-pollinating. The maximum is 48.9% under
Ma. They show 36.1 under red and 54.9 under white, or
4 and 12.1 more than the percentages of those colors.
Of the non-pollinating visits 80% are to red. These show
percentages of Lepidoptera species and visits: Hesperi
41.5, 44.6; Papilion 13.2, 18.7; Nymphal 9.4; 12.7; Pier
9.4, 12.4. These families show maxima under Ma, The
Lycaen show maxima under Mis.
Coleoptera and Hemiptera.—Are 6.8% of the visitors
and make 3.5% of the visits. The maximum of 48.3% is
under Pol. They show 12.9% under red and 64.5 under
white, or 19.2 less and 21.7 more than the percentages
of those colors,
Early visits at Inverness and Carlinville—Comparing
Percentages of the visits observed at Inverness, Ja.-Ap.,
with those observed at Carlinville before July shows some
remarkable results. The Lepidoptera are the most con-
Spicuous with 29 against 6.4. The lower Hymenoptera
show 21.4 and 10.1, resembling the late visits at Carlin-
ville, 22.7. ‘The lower Hymenoptera and Lepidoptera
324 Trans, Acad. Sci. of St. Louis
make 51.7% of the early visits at Inverness and only
16.5% of those at Carlinville.
The short-tongued bees show 12.4 and 27.7, while the
long-tongued show 13.0 and 20.7. The bees make 48.5%
of the early visits at Carlinville and 25.4% of those at
Inverness. The short-tongued bees and flies make 56.9%
of the early visits at Carlinville and 32. is of those at
Inverness.
BIBiLioGRAPHY.
1. Robertson, C. 1887 (12). Fert. Calopogon. Bot.
G. 12 :288-91.
2. ——,1888. Zygomorphy. Bot. G. 13:146-51, 203-8,
224-30.
3. ——, 1889 (5). Fis. I. Bot. G. 14:120-6:
4. ——, 1890. Fis. Umbelliferae. St L. A. 5:449-50.
5. ——, 1891 (2). Fis. Asclep.Scroph. St. L. A.
5 :569-98.
6. ——, 1891 (3). Fis. VI. Bot. G. 16:65-71.
7. ——, 1893 (2). Fis. X. Bot. G. 18 :47-54.
8. ——, 1923 (10). Visits of Insects. Psyche 30 :158-69.
9. ——, 1924 (9). Fls. XXIII. Bot. G. 78 :68.84.
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Transactions of the Academy of Science of St. Louis
a
“ps
ieee AA Se rongpe
Volume XXV, No. 9
FIELD STUDIES IN THE BEHAVIOR OF THE
NON-SOCIAL WASPS
PHIL RAU
FIELD STUDIES IN THE BEHAVIOR
OF THE NON-SOCIAL WASPS
CHAPTER
CHAPTER
CHAPTER
CHAPTER
CHAPTER
CHAPTER
Co A ™™ER
~ te,
xy.
VE.
VOT.
PHIL RAU
CONTENTS
The life and habits of two beetle-hunting
wasps.
Cerceris raui
Cerceris bicornuta
An amputator of spiders’ legs.
Pseudagenia mellipes
The grass-carrier wasp.
Chlorion auripes
An assortment of twig dwellers.
Diphlebus biparitor
Diphlebus tenax
Silaon niger
Solenius interruptus
Stigmus fraternus raui.
Hypocrabro stirpicolus
Wasps of the genus Odynerus and their
nesting habits.
Odynerus (Stenodynerus) conformis...
Odynerus (Stenodynerus) pennsy wk
POT TT ea eh ee eerie Soy cea
Odyneris Leucomme lars cceescecseceveenveeeeneneen ate
Odynerus foraminatus ... saith
Odynerus capra
The patriarchate wasps of the genus
Trypoxylon,
Try posylon Clavat urn a neenenn mmm
Trypoxylon politum Seen
Trypoxylon sates Ee ag ee
Trypoxylon texense .. Rigo SR
The nesting habits of the ae
mud-dauber
Sceliphron caementari
The prey of Sctliphows: ibaa: =
PAGE
325
337
INTRODUCTION,
This work on the behavior of twenty-three species of
solitary wasps was all done in the vicinity of St. Louis,
Mo., between the years 1917 and 1923.
As in previous studies, all insect material was sub-
mitted to experts for identification, and their names ap-
pear in brackets throughout the work. The photographs
Were made by the author, and the drawings were made by
Dr. Gustave Dahms of this eity. A debt of gratitude is
due to my wife for valuable assistance on the manu-
Script.
‘Observation sets the problem; experiment solves it,
always presuming that it can be solved, or at least if
powerless to yield the full light of truth, it sheds a cer-
tain gleam over the edges of the impenetrable cloud.’’
—J. H. FABRE,
“So the first task of comparative psychology is to find
out the instinctive equipment of any animal studied. In-
stincts are, however, well worth study for their own sake.
An instinctive fear of a certain enemy may be as truly
useful to an animal as sharp teeth or protective colora-
tion. Instincts are the expression of structures and func-
tions of the nervous system, and are as real and as im-
portant matters for the biologist as are bones and blood
vessels.’’
—E, L, THORNDIKE.
‘It may be noted that all the activities of ants, their
reflexes and instincts as well as their plastic behavior,
gain in precision with repetition. In other words, all
their activities may be secondarily mechanized to form
habits, in the restricted sense of the word. This is tanta-
mount to say that even the reflexes and instincts are not
So stereotyped but that they may become more so by ex-
ercise during the lifetime of the individual. And not only
do ants thus form habits, but . . . these habits when
once formed are often hard to break. It is certain that
many instincts . . . are at first complete and indefi-
nite and are guided to their proper course by stimuli
that affects the organism at a later period. .... There
is little doubt, moreover, that the more fixed or stereo-
typed instincts are phylogenetically the older.’’
—wmM. M. WHEELER.
Sa Nats
=
Field Studies of the Non-Social Wasps 325
CHAPTER I.
Tue Lire Hasirs or Two BEEtLe-HUNTING WASPS.
THe Jerseypate Cercents, Cerceris raui Roh.
[S. A. Rohwer].
Jerseydale (fig. 26), some 30 miles south of St. Louis,
consists of a railroad sign-post, a milk-can platform, and
a well of excellent cold water; this equipment renders it
easily recognizable as the center of a thinly populated
but prosperous dairying community.
Besides the well, Jerseydale has other attractions, un-
noticed perhaps by the men who daily drive up to the
platform to exchange their full cans for empty ones. In
the hard road under the hoofs and wheels abound the
Cerceris wasps, Cerceris raui. This wasp (fig. 27) was
described by Rohwer from material submitted from three
distinct localities: Lake View, Kan.; Wickes, Mo., and
Jerseydale, Mo.* While only fragmentary notes were
gathered from the first two localities, at Jerseydale the
life secrets of this species were more fully discovered.
One salient feature of their habits is that in all three
places their nests were dug chiefly in the hard-packed
roads, where traffic was heavy, and only a very few nests
on the outskirts of a colony occurred at the roadside.
The hole which this species makes is an excellent piece
of workmanship. There is no attempt at concealment of
the burrow, such as we find practiced, sometimes very
skillfully, by some other species of wasps. Her nest is
So inaccessible in its situation that she can well afford
to work boldly and openly before all men, without fear
of her home being molested, unless by parasites which
are cunning enough to elude her and gain access by her
own doorway. The burrows in each of the three habitats
*Also taken later at Wesco, Mo., 108 miles west of St. Louis.
326 Trans. Acad. Sci. of St. Louts
were found in the middle of a well-beaten road, where
the earth was so hard packed as to be almost impenetra-
ble with ordinary tools. ‘A spading-fork, a digging-trowel
and a pruning-knife each in turn proved ineffectual in
the conquest, and only by the aid of a hatchet was I
finally able to follow a very few burrows to their termini.
The work was rendered increasingly difficult by the fact
that the nests were in the beaten track, and at the ap-
proach of each automobile or wagon it was necessary for
the perspiring investigator to take flight to a safe dis-
tance to escape the wrath of the irate drivers who seemed
not to appreciate the beauty of the quest of truth when
it led to the digging of chuck-holes in their roadway. At
Lake View the road leading to a country club was beaten
so hard by the automobile traffic that I could not dig up a
single nest. At Jerseydale the yellow clay was very
sticky when wet and flinty when dry, and at Wicks (fig.
28) the admixture of clay and rocks rendered digging
most difficult. At the two latter places, however, a few
nests were found at the side of the road which could be
dug out to sufficient depths to learn some details of their
interiors.
The mouth of the hole is always beautiful in its sym-
metry; it is 14 inch in diameter, and around the aperture
is a mound of chips or granules of earth which have been
brought up from below (see fig. 29; a ten-cent coin nearby
shows the relative size). The burrow is amply large;
the diameter is sufficient to permit the occupant easily to
turn around or even to make her toilet while within. The
tunnel is long and tortuous. In the places where rocks
are present in the soil this is, of course, the logical re-
sult, but the same condition exists to an extreme degree
in clear soils also. The illustration (fig. 30, %4 nat. size)
gives an idea of the rambling course of one of these chan-
Field Studies of the Non-Social Wasps 327
nels, but, of course, an incomplete idea, since a diagram
can show the curves in only one plane, and not the curves
toward and away from the observer. Another was de-
scribed in detail in my notes as follows: This hole went
straight down for 2 inches, then curved toward the south-
east for 3 inches, then southwest for 4 inches, made a
sharp turn toward the northwest and downward for 4
inches, westward and still down for 2 inches, upward and
west 2 inches, then curving downward and south about 4
inches. Along this section of the tunnel were three cells
containing fat Cerceris larvae and beetle remains. These
cells were oval and about 114 inches long and % inch in
diameter. Beyond this the tunnel continued slightly up-
ward and westward for 3 inches more, and in the end,
in a sort of pocket, were 3 more fat beetles. This indi-
cates that the wasp digs her tunnel and makes several
pockets at the terminus for brood cells. The tunnel
varied slightly in diameter, the minimum being about %
inch. The total length of this one was 24 inches, and the
cells were situated 17 inches below the surface of the
ground. One can readily see what an enormous task this
mother wasp had accomplished, to have made so large an
excavation in soil which was so nearly impenetrable that
I myself could hardly dig it.
The excavation of another burrow was begun in the
absence of the mother wasp. The hole was likewise very
tortuous, and at a depth of 15 inches it turned directly
under the hardest part of the road, where the tools broke
and the digging had to be abandoned. Meanwhile the
mother returned, and so persistent was she in trying to
enter the wreck of her home that twice I picked her up
in the forceps with perfect ease. She fairly fought to
get back to the place, and dug for more than an hour
trying to follow her old burrow, which she recognized so
328 Trans. Acad, Sci. of St. Louis
surely despite its mutilation. I left her still working
feverishly at her task of reconstruction.
Since in the three places where these wasps have been
found they occurred in settlements or colonies, one won-
ders if they do not dig their way out of their subter-
ranean winter dormitory in the spring and then turn
around and enlarge their exit-passage for their new bur-
rows. It seems incredible that one wasp is able to dig so
large a tunnel in so short a time.
I earnestly hope that the reader will be satisfied with
the details of these few burrows, so I may mercifully be
spared the task of digging out another one. So far as I
could learn from others partly excavated, the tortuous
course and the dimensions of these are fairly typical.
I have not been so fortunate as to see the beginning
of the digging of the burrow. Later on in the process,
however, when the tunnel was only being enlarged or ex-
tended, I have observed the outward evidence of work
going on within. There is then little display to attract
the attention of the passer-by; the wasp in her dark tun-
nel down under the earth digs up the soil—I know not
how—and packing a considerable mass behind her body
and above her in the tunnel, backs up and pushes it out
above the surface with her abdomen. This soil comes
out, not in loose, powdery form, but in granules
packed together to form plugs or sausage-like masses,
which protrude for a time above the surface and later,
when thoroughly dry, collapse in a heap at a touch or
breeze. This digging was observed several times early
in the day, before the wasps had come out of their holes
to brush back the dirt from around the mouth of the
burrows, and depart for refreshments or prey. Hence
I cannot say with fairness that these Cerceris are not
early risers, for I do not know how early they begin their
Field Studies of the Non-Social Wasps 329
work in the dark gallery, but they do not open up their
doors, which are closed each night, until the sun is high
and hot in the summer sky. This statement implies that
they sleep in their burrows, and so they do; at least the
females of the population which possess burrows sleep
there; I have not yet been able to find, by careful scrutiny
of the vicinity, where the males find their night’s lodging.
I have never yet seen the males enter the burrows,
although they are frequently to be seen pursuing the fe-
males as they come and go. They are small and agile,
and difficult to apprehend. The sexes are so different in
appearance that they look like two different species, the
female a large, reddish-colored insect, and the other
much smaller and yellow. Mating occurs on the wing or
while the home-maker goes to and fro about her work;
no special frolic or festival has been observed to accom-
pany this function, such as occurs in several species of
wasps.
The owners of the burrows come home and enter the
nest early, about 3:30 to 4:30 o’clock, and push up a
plug of loose earth to close their door snugly for the
night. This roof remains in position remarkably well,
in spite of the fact that it is so soft that it will collapse
if tickled with a straw. On one afternoon when a shower
occurred early, I visited the colony and was surprised to
find that even at that time of day, which is usually the
wasps’ busy hour, these good housewives had all hurried
home to shut up the house before the storm broke; every
door of the fifty or sixty nests was closed from within!
In covering the hole the wasp backs up with the ab-
domen extended straight and the load of earth under
the abdomen and behind the legs. Thus she pushes the
load up to a point 14 inch from the top, where she presses
the dirt against one side of the hole. Thus she brings up a
330 Trans. Acad. Sci. of St. Louts
few more loads, condensing and packing the earth against
one wall; then she brings up more dirt to close the re-
maining aperture. Even then she does not stop, but adds
many layers to the plug from the inside; we are made
aware of this by the movement of the plug as it heaves up
with the vigorous pressure from underneath as each load
is packed in place. In many cases the wasps seem to
deem their day’s work done at 4 o’clock, and by 6 most
of the holes are partly or completely closed for the night.
It is a pretty sight in the morning to see the wasp first
venturing out. The granules of clay covering the hole
quiver and part, the head appears, and the opening is
silently made wider. Then the wasp pauses to peer cau-
tiously about (fig. 31); this habit of cautious recon-
naissance is a very characteristic gesture of this species.
She creeps so stealthily to the edge of the hole that one
is hardly aware of the motion, and pauses with her face
at the aperture and peers about with a gaze so intent
that one can almost feel it; if she is not satisfied that all
is safe, she drops stealthily back; if all is well, she darts
away like a flash, leaving the burrow wide open. She has
retreated thus when I moved noiselessly at a distance of
4 feet; this would indicate that her range of vision en-
compasses at least that distance. Upon returning home,
she seems to have no trouble in locating her burrow, al-
though it is one among so many all alike, in the roadway
or in the sparse grasses nearby. She does not loiter,
but plunges headlong into the burrow, almost before one
has time to get a good view of her. Sometimes she circles
about on the wing a few times before she seems to get
her bearings to make an accurate descent directly upon
her burrow.
Every observation upon the Cerceris raui population
at Jerseydale indicated that this wasp uses only two
Field Studies of the Non-Social Wasps 331
species of Coleoptera as food for her young, viz., a short,
fat weevil, Thecesternus humeralis Say [E. A. Schwarz],
and the long, slim weevils, Liaus concavus Say [B, A.
Schwarz] (fig. 32). Less extensive observations at Lake
View, Kan., and Wickes, Mo., gave the same results, but
the data there were not sufficient to be conclusive. Of 22
females which were caught as they returned to the nest,
16 carried Thecesternus humeralis, and 6 had the long,
_ Slender beetles, Lixus concavus. In the several brood-
cells which I had the good fortune to explore, the former
beetles were present in far greater numbers. In some
cells all the provision was of that species. About 15 to
22 beetles seems to be the ration allowed each young
wasp.
In all of the cells wherein the larva had already
worked over the food supply provided for it, we found
that it had separated each beetle into the three com-
ponent parts, the head, thorax and abdomen. The head
separates easily from the thorax in these beetles, and the
larva undoubtedly takes its first meal from the soft tis-
Sues at the point where the thorax joins the abdomen.
It is wonderful to see how clean these beetle shells have
been licked when the larva is done with them. (Fig. 33
Shows the beetle remains after the feast). One wonders
how it is possible for the larva to clean out the contents
So completely without breaking through the body wall
at various places, but when one sees the remarkable
adaptation of its anatomy for just this purpose, one can
readily see how it succeeds in reaching all parts of the
body cavity. While it in general looks like all wasp larva,
the oral end tapers into a nice point so that it is possible
for this part to enter the abdomen through one opening
and reach clear to the other end. The head part sways
332 Trans. Acad, Sci. of St. Louts
from side to side, and the jaws move about seeking the
food when it is taken from them.
The beetles, when they are removed from the cells, are
motionless, but if they are taken from the mother wasp
as she brings them in, and placed in a vial, they soon re-
vive so that they climb over one another very actively.
Hence one doubts if they have actually been stung. The
22 taken from the wasp mothers, as previously stated,
were very active when examined the same day; when in-
spected two days later there was much excrement in the
vials, indicating that life had continued for some time,
but the beetles were dead; whether death had been due
to the confinement or the sting, I do not know. Close
observation has taught me that these wasps do sometimes
sting their prey, but how general this habit is has not
been ascertained. One of them stung a live weevil which
{ substituted for her own, while another did not. When
the wasp was out foraging I placed a small ball of cotton
in the mouth of her burrow; when she returned and at-
tempted to enter, she dropped her prey in alarm and
flew away, and when she returned she found I had re-
moved the plug. Meanwhile I had also exchanged her
quiet weevil for a live one, which was so active that I
could hardly keep it near the hole. The mother returned,
flew directly into the hole as usual, and after a few sec-
onds crept to the top, poked her head out cautiously,
grasped the beetle in her jaws and dragged it in; if a
sting was administered it must have been done under-
ground.
Many wasps, in foraging or bringing in building mate-
rial, take a direct flight to and from the place of interest,
either with or without circles of orientation. This species,
so far as I have observed, follows a rather strange
course; she does not fly ‘‘as a bird flies,’’ or ‘‘in a bee
Field Studies of the Non-Social Wasps 333
line,’’ but she literally makes a round trip. Several which
I watched going to the cornfield on the other side of the
railroad track, left the nest flying northward; after fifty
feet or so they circled around toward the east, and after
an absence of from 3 to 8 minutes they came back with
their prey, by way of the south. Just how general this
habit of circular flight is I do not know.
They work with surprising rapidity in bringing in their
victims. I timed a number of them on different occasions
and they usually took only 3 to 8 minutes to capture a
beetle and return. They lost no time in finding the open
nest, plunged in, and stored the booty speedily.
It was noticed that practically all of the weevils
brought in by these Cerceris were covered with a crust of
earth. That raises the question: do these beetles oceupy
subterranean habitations where these wasps are obliged
to dig for them? The wasps seem to find them so easily
and in such large numbers that there must be an abun-
dant supply of them near, perhaps in the cornfield. To
ascertain the hunting habits of these wasps one ought
to know something of the two species of beetles that they
hunt, Livus concavus, properly known as the rhubarb
cureulio, and Thecesternus humeralis. Of the latter spe-
cies, Mr. E, A. Schwarz writes: ‘‘The life history of
this weevil still remains unknown, but the imago is quite
common under dried cow dung in our prairie states, or
under stones in dry situations in Indiana, western Mis-
souri, Nebraska, Colorado, New Mexico and Texas. Most
of the specimens in our collection are covered with a
thick crust of dirt, which shows that the earlier stages
are passed underground and that the larvae will be found
In the roots of some plants. Oceasionally specimens of
the imago have been found above ground on the stems
of various plants. * * * A somewhat allied species,
334 Trans. Acad, Sci. of St. Lows
Rhigopridius tucumanus Heller, from Argentina, has
been bred from the tuber of a potato.’’
The weevil makes so small a burden that the Cerceris
can carry it very easily and swiftly. In all of the cases
which I could observe closely, the wasp carried the prey,
clasped in her middle legs, with the ventral side of the
weevil against the under side of her own body, and in
some cases it appeared that she grasped its proboscis
in her jaws. She drops her burden quickly if disturbed.
One wasp returned with her weevil under her body, and
as she found difficulty in finding her hole, which had ac-
cidentally been covered, I pressed close to see her method
of search. She braced herself with her hind feet while
she dug in the mound of dust with her free fore-feet.
Not gaining much headway in this manner, she dropped
her weevil and then worked faster with four feet. Thus
she eventually found her hole on the other side of the
mound. Meanwhile, I exchanged the weevil which she
had laid down, for one borrowed from another wasp.
Several times she went into the hole, as if to make sure
that everything was right, and came out again. At last
she approached the weevil, took it in her mouth, rolled
over and stung it. In my eagerness I came too close and
frightened her away. After ten minutes more of nervous
coming and going, she again took the weevil in her mouth,
curled her body almost into a circle in order to reach the
prey with her sting, propping herself up meanwhile on
her two protruding wings, while she inserted her sting
on the ventral side of the victim and kept it there for
almost a minute, her abdomen pulsating all the time. All
this happened on the mound of loose soil, only a half
inch from her hole, which was now open; nevertheless
she took up her weevil, flew into the air, circled about a
Field Studies of the Non-Socral Wasps 335
few times, alighted again in the normal manner and
plunged in.
While another nest was being observed, a troublesome
weed that grew near it was pulled up; this changed the
topography of this tiny area. Upon returning several
hours later we found four weevils lying out on top of the
ground, and the returning wasp even then circled and
buzzed about the hole for fully half an hour before she
would enter. It was evident that she had discarded the
weevils lying there in the sun; either she had dropped
them as she had come back from foraging and had lost
them in her confusion at finding the familiar landmarks
altered, or she had suspected that, since the region had
been changed, her nest had probably been meddled with
and so, like other wasps of our acquaintance, she had
thrown out the stores she already had as though they
were contaminated.
One nest was especially conspicuous because at that
point in the road the surface of the ground was covered
with black cinders, while the circular mound of subsoil
around the hole was gray. This contrast made the nest
itself a conspicuous landmark for the returning wasp.
I watched until the wasp left home; then with the point
of the trowel IT carefully removed this gray earth and
replaced it with a mound of black cinders, just like the
surrounding medium. I was quite proud of my new ac-
complishment of building wasps’ nests, for the imitation
was, to my eye, very accurate except for the color of
the material. It was, of necessity, a hasty job, for the
rightful proprietor returned from her trip in only two
minutes. She flew direct to the location of her hole as
usual, but just above it she brought herself up with a
Jerk and flew to and fro over it in an agitated manner
for several minutes. Her confusion was evident; twice
336 Trans. Acad. Sci. of St. Louis
she dipped down on the wing as if ready to enter, then
withdrew in alarm; again she tried to scrape away the
cinders a little to one side, as if thinking that her burrow
had been covered by accident, as frequently happens dur-
ing the mother’s absence. Thus she continued to search
in all the crevices and possible places. For the first five
minutes of this hunt, she carried her prey with her; then
she dropped it until she could find her burrow. After that
she searched on foot thoroughly over all the area within a
dozen inches of her hole, only occasionally wandering in
circular flights over a wider area. She seemed sure of the
general region—the cinder area—but the ring of gray
dust around her hole was not to be found; therefore she
did not recognize it as her own even after poking her head
in three times, and went back to scratch among the cinders
in various points for another quarter of an hour. After
she had spent a full half hour in this faithful search, I
tried to make reparation by bringing a trowelful of gray
dust of just the right shade and fineness and putting it
nicely around the hole. As soon as I withdrew she re-
turned on the Wing to continue the search, and as she came
within sight of the place, she dashed straight to it and
tumbled in headlong, in the old familiar manner! She
remained in longer than usual this time, then passed out
and in several times, making short flights of orientation
about the place each time. Her faith had been severely
shaken and she had to reassure herself many times. Fur-
thermore, things were not yet just as they should be; my
clumsy hand had spilled some cinders in her burrow, and
she had to make the necessary corrections and get ac
quainted with the place anew. At last, with one more
careful flight of orientation, she departed for the fields to
resume her foraging.
These wasps keep so diligently to their nesting business
Field Studies of the Non-Social Wasps 337
that I have found little else to record concerning them.
They neither loiter nor dance, fraternize nor fight. Only
a few times during the summer were they seen feeding
quietly on the flowers of buckbrush and goldenrod near
their nests.
The wasps continued to work the old burrows until
late in the summer, but no new ones appeared during the
latter part of the season. They continued to push up dirt
and carry in prey; hence I surmised that they were ex-
tending the burrows and putting in additional pockets.
During the week of September 12-18 they disappeared ;
the earth over their holes was soon beaten down by rain
and traffic. The place and manner of their death was not
learned.
A number of other species were seen prowling about
the premises of Cerceris’ home. Their purposes or meth-
ods of approach were not actually ascertained, but we
know the character of some of them well enough to feel
justified in holding our suspicions. A Parametopia sp.
entered one hole and was captured as it emerged. In
early August many Hedychrum violaceum Brulle [S. A.
Rohwer] (fig. 34) were seen entering the burrows, and
in two instances Lyroda sublita Say [S. A. Rohwer] in-
truded. A Megachile was also seen hovering about the
burrows of Cerceris, but I could not catch her in any mis-
demeanor. Several specimens of the Dipteron, Exopro-
Sopa fasciata Macq. [C. T. Greene] were taken as they
persistently hovered above the Cerceris burrow at Jer-
Seydale,
Tue Bu-sue Huntress, Cerceris bicornuta Gurrty
[S. A. Rohwer].
Cerceris bicornuta, like C. raui, digs holes in the earth
and uses bill-bug beetles for food. The various species
338 Trans. Acad. Sci. of St. Louis
of Cerceris do not all have the same habits; there is much
diversity within the genus, in both nesting sites and prey
eaptured. While most members of the genus nest in the
ground, we have a few that make nests in twigs, e. g., C.
finitima Cress., and in getting prey for the young, certain
species catch weevils, Buprestid and other beetles, while
others seek bees.
If there are such things as plant societies, then surely
we may say insect societies exist. When one compares
the wasp population of our baseball diamond at St. Louis
with that of the school yard at Pevely, thirty miles dis-
tant, then one must surely say that certain types of areas
attract certain insect societies. The two places gave us
Bembia nubilipennis, Odynerus dorsale, Cerceris bicor-
nuta, and one single specimen at each place of Cerceris
fwmipennis. The proportion of each species in the popu-
lation as a whole was likewise similar.
One frequently finds the hole of this wasp closed when
the digging is still in progress, but it is easily located and
partially identified by a pile of loose dirt over it (fig.
35), at the center of which is usually a group of a half
dozen rolls of dust, tightly packed, just as the wasp has
pushed it out of the burrow. This detail indicates that
her method of excavating is similar to that of C. raw.
She digs and pushes the dirt up to the upper part of the
channel where it forms a plug; when at intervals she
pushes this whole stopper out above the ground the
masses of dirt retain their sausage-like form for a time
after they have been forced out, until they become thor-
oughly dry and crumble to dust in the wind. Certain
beetles excavate in the same way. It seems that excavat-
ing and provisioning must be carried on at the same time,
for often when she pushes out a pack of dirt, she throws
out some of the bill-bugs as well. Four beetles were
Field Studies of the Non-Social Wasps 339
found in the dust on the top of one of these nests. These
weevils ‘‘play possum,’’ and one cannot always tell
whether they are dead from the sting or merely in a feint,
but these showed no indication of life, even when they
were pinned without the usual formalities of a cyanide
bottle. :
These wasps dig with surprising rapidity. In one nest
where the loose dust on the surface was brushed away at
7 o’clock one evening, the next morning at 9 there was a
mound of earth 2 inches in diameter and 34 inch high
which had been newly thrown out of the burrow. It
Seemed impossible that this mound could have been
thrown up since daylight; hence I suspect that the mother
Verceris had been at work during the night. After all,
how can they tell day from night when they are at work
in the inky darkness of their underground tunnels?
Rain works real hardship to these little creatures. One
of my records tells of a nest in which the work was pro-
gressing nicely, and each morning showed a mound of
fresh dirt on the surface. A steady rain soon flattened
the mound, but the plug held for several hours; a down-
pour eventually washed it down, however, and the hole
was flooded. The next day I watched the nest and con-
cluded that the owner had been drowned, for during sev-
eral hours of sunshine she did not appear. In the middle
of the afternoon, however, I saw her again carrying in
her eatch of bill-bugs! How she could have escaped death
by drowning I cannot see. In other nests which we
watched, however, the plugs were soon washed down;
the occupants of most of these nests never reappeared.
Three survivors tried to rebuild their homes, but were
defeated by heavy rains. Thus we see that the rain here
was a factor in exterminating the adult population; how
the young fared in these circumstances we do not know.
340 Trans. Acad, Scr. of St. Louts
‘After the excavating is once done, the workers seem
to have no objection to leaving the mouth of the burrow
open. They are frequently to be seen on sunny days,
with the face just within the aperture of the hole, their
bright eyes shining as they gaze into the outer world.
But if one comes too near, they, like C. raui, creep down,
cautiously, almost imperceptibly; one does not realize
that they are moving until they have vanished from
sight.
While Cerceris bicornuta persistently chooses for her
nesting site a hard-packed soil, yet a number of times I
have found evidence that she gladly avails herself of
assistance in her digging by utilizing some hole already
begun. In one case, one extended a neat hole left by a
peg which had been driven into the ground and pulled
out. In other instances they used and lengthened the old
holes from which Bembiz nubilipennis had emerged (fig.
36). While they do not seem actually to follow up the
Bembix or depend upon them for their assistance, they
frequently have the opportunity of availing themselves
of it, since their choice of the same environment often
brings them together.
When one finds insects established in a city lot, one
is inclined to think of them as relies from a preciviliza-
tion period, clinging to their former habitat despite the
tightening about them of urban conditions; one is sur-
prised to learn that species new to the neighborhood are
coming in and becoming established under our very eyes-
Intensive Hymenoptera collecting in this vicinity for
eleven years had not revealed one specimen of this wasP-
The small area where they are now established was dur-
ing the four years, 1913 to 1916, subjected to my very
intensive study; observations were made almost every¥
day as I crossed the field, yet I never found a single speci-
Field Studies of the Non-Social Wasps 341
men of C. bicornuta. In 1917 I was out of the city, and
the next summer, on June 30, the first specimen was cap-
tured; during the following week six others were found
nesting there. This shows how quickly and firmly they
become established when they find a desirable environ-
ment.
C. bicornuta preys upon three species of bill-bugs,
Sphenophorus placidus, S. zeae and S. parvulus. Bill-
bugs are very destructive to cereal and forage crops, and
the Cerceris wasps are one of the important natural
checks. The blue-green bill-bug, 8. parvulus, is a widely
distributed upland species usually infesting timothy and
bluegrass, but often injuring wheat, oats, barley and rye.
Pupation occurs either in the corms or in the soil.
A large blue cuckoo-bee, Hedychrum violacewm, en-
tered the burrow of this wasp and remained within for
twenty minutes; what her errand was we can only sur-
mise,
Adults of this species have been found feeding on the
flowers of the buckbrush.
342 Trans. Acad, Sci. of St. Louis
CHAPTER II.
Aw Amputator oF Sprpers’ Leas. Pseudagenia mellipes
Say [S. A. Rohwer].
The genus Pseudagenia belongs to the family Psam-
mocharidae, whose habits of nesting are in rather un-
stable conditions, since each species shows peculiarities
of its own. Natural selection has not, up to the present
date, created fixed habits for the genus. There is, how-
ever, one department in the work of this genus that has
become rigidly established, that is the cutting off of the
legs of the spiders stored as food for the young. All
species whose habits have been studied show this trait,
although in P. mellipes (fig. 37) it is possibly not so rigid
as in other species, for in numerous cases I have found
that only a part of a spider’s legs have been removed.
As in the genus Trypoxylon, we find great versatility
of behavior within certain species and within the genus.
To Francis X. Williams* falls the credit of having
brought to light some highly interesting behavior infor-
mation on seven species of Pseudagenia which were stud-
ied in the Philippine Islands, with the aid of which one
can point out the evolution of the nesting behavior, from
the simple, single mud cells of certain species to that
masterpiece of economy, the work of P. mellipes.
Versatility in behavior, or adaptability, seem to be the
qualities which have led some individuals, at least, of
P. mellipes to a method of nesting that is most economi-
cal of labor, materials and time. In connection with the
evolution of the nesting habits of Pseudagenia, the fol-
lowing statement by Williams is indeed significant: ‘* The
“Philippine Wasp Studies. Rept. Exp. Sta. Hawaiian Sugar Plant.
Assoc., Bull. 14, pp. 79-108, 1919.
Field Studies of the Non-Social Wasps 343
genera Macromeris, Paragenia, and Pseudagenia and
others [of the Psammocharidae] are mason wasps, hav-
ing advanced beyond the digging stage still adhered to
by the majority of the family. They build cells of clay
or other earth-like material; they may construct these in
sheltered or unsheltered places above the ground, more
rarely in burrows.’’ As Williams says, ‘‘having advanced
beyond the digging stage,’’ to a mud-daubing cell-maker,
So certain individuals of the species P. mellipes, in their
cell-making, have gone a step forward in eliminating the
carrying of the building materials. We find that in P.
mellipes this labor-saving method of using materials at
hand, or rather of finding the building materials in the
Shape of a lump of mud plastered on the wall, and then
fashioning it into cells, entails the habit of carrying
water. This method, when first I discovered it, greatly
surprised me, since I thought only the Odynerus wasps,
and Anthophora bees were capable of so ingenious a
method. Since then I have found this same feat done
by Chalybion caeruleum, which shows at least that this
habit can penetrate families regardless of structure (tax-
onomy). That this water-carrying habit is more deep-
Seated in the genus than the newly acquired habit of
using mud nests already at hand is evidenced by the fact
that Williams tells us of a species in the Philippine
Islands which follows this method, showing that habit
persists in spite of remoteness. Another habit that dis-
tance has not obliterated is the method of using the
dorsal tip of the abdomen, flexed under the head, as a
rounded smoothing tool, in fashioning the nest. This
habit we observed also in Pompiliodes tropicus,* and in
Williams’ report he records this habit for three species
of Pseudagenia,
“Wasp Studies Afield, p. 54, 1918.
344 Trans. Acad. Sci. of St. Louis
The stability of these two habits, in groups of or-
ganisms separated from each other by half the girth of
the earth, lends a tinge of romance to the subject. Shall
we say that these habits are instinctive and of so strong
a type that they persist regardless of time and space,
or shall we rather say that these groups of insects, m
the two remote regions, have independently acquired the
same habits, through the similarity of their problems or
through the similarity of their environment?
I said that in the evolution of nest-building activities,
P. mellipes is in advance of any other member of the
Pseudagenia clan, despite the fact that the Philippine
Pseudagenia are likewise water-carriers and likewise use
the abdomen as a tool wherewith to fashion the nest. The
facts are that the oriental wasps still carry the building
material to the site, while some individuals of our P.
mellipes have advanced to the point where the carrying
proclivities are eliminated. Not all of our P. mellipes
have abandoned their habit of carrying clay, for we have
formerly recorded three and four-celled nests taken from
under loose pieces of bark (fig. 40), and a four-celled nest
that was found in an oak-apple still hanging on the tree.
All of these nests in due season gave forth P. mellipes,
and both situations certainly entailed the carrying of mud
from a distance. Other nests which must have involved
the new method of building, have been found in old nests
of the pipe-organ wasp, Trypowylon politum (fig. 63),
and in the abandoned nests of Sceliphron caementarium
(fig. 38), where these wasps had built their little cells
within the larger ones of both species. Thus we see the
most recent accomplishment of the species, that of carTy-
ing water, to be so new as not to have permeated the race,
and the pioneers in this habit are the wasps observed at
an obscure corner of the earth called Wickes.
Field Studies of the Non-Nocial Wasps 345
It is fascinating indeed to think of our own little hand-
ful of P. mellipes as displaying perhaps a more advanced
point in their evolution than other members of even their
own species, and exhibiting a type of behavior more ad-
vanced than that of any other member of the genus whose
habits have been recorded. In this connection we must
not overlook one Philippine species, P. nyemitawa, whose
unique habits are recorded by Williams. This wasp
pastes its nest, consisting of two or three cells, on tree
trunks, in rather exposed situations, but the structure
seems proof against wind and weather, being varnished
over with a tree gum. The cells are made of an earth-
like substance such as is used by the termites for their
Coverways. ‘‘The wasps first sip up some water from
Some convenient hollow or edge of stream, and the ball
of earth subsequently gathered is worked around in her
mouth until it assumes the right consistency, when it is
plastered on the building site with the dorsal tip of the
abdomen. * * * Before the cell group of two or three
cells is completed a partial coat of varnish may be put
on. The cells finished are more closely united with mud.
x When Pseudagenia is through with the mud she
turns her attention to a tree gum which she works over
in her mouth-parts and spreads on precisely as she did
the mud. Then without intermission she brings in an-
other final coating or rather patching material—a pale
8tay lichen—which she works up in her mouth into a
Sort of viscid paste also applied like mud and varnish,
but the nest is blotched rather than completely covered.”
There is a small outbuilding at Wickes which harbored
a number of old mud nests of Sceliphron caementarium
in addition to a few nests of Polistes pallipes. For two
years, 1917 and 1918 the inhabitants therein were ob-
jects of study. The ecological succession of life, even in
346 Trans. Acad. Sci. of St. Louis
so small a room, was as clear as it was interesting. To
recapitulate the story of the occupants, the original
builders, Sceliphron, had made their nests and left them
to the insect community; then the succeeding inhabitants
followed in peaceful succession. First after the Sceli-
phron came renters by the name of Trypoxylon clava-
tum. These were abundant, and used the old mud nests
with only slight modification. After T. clavatwm had
been watched and studied here for two years, it was
suddenly discovered, when the season of 1919 opened,
that clavatum was no more; that a species never here-
tofore seen in this building was present in fair numbers,
and had taken possession of the situation for its abode.
This new citizen in the community, that had the art of
making dainty little nests out of old tumble-down, twice-
used mud huts, was Pseudagenia mellipes.
In previous descriptions and in the paragraphs preced-
ing I have already shown that P. mellipes is a fairly
versatile little creature. In a former work, when we re-
corded that this insect was found in the old cells of Scelt-
phron (fig. 38 shows the old nests broken to expose to
view the mud cells of P. mellipes), and discovered heavy
mandible marks on the outside of the nest, we registered
the suspicion that this was the work of P. mellipes. This
suspicion was confirmed by the observation of this lot
of little creatures in action. The following extracts from
my notebook will give the details.
At 12:50, on September 6, I noticed a one-celled nest
of Sceliphron with a P. mellipes aboard. The opening
was half sealed in a manner which was new to me; @
wall was being built across the aperture from the two
opposite sides, leaving a vertical crack across the center
about 5 mm. wide. After five minutes the opening Was
smaller. My chief attention was at that time concel-
Field Studies of the Non-Social Wasps 347
trated upon other insects; when again at liberty, twenty
minutes later, I found the cell completely sealed, and then
I ruefully realized that I had lost a rare opportunity to
observe this species in action; to find whether she got the
mud from elsewhere to seal the aperture, or in some way
worked out the mud from various parts of the nests. For
the next hour I watched the wasp busy at her work on
various parts of the cell, principally at the newly made
plug, in a very interesting manner. With her jaws she
would scrape very energetically about the rough wall of
the shanty, trying apparently to remove some of the sur-
face of the wood, but in reality she was scraping together
whatever she could get (often it was mostly dust and spi-
der web) which she worked in her jaws on her way back to
the nest a few inches away. When she reached the nest
the mouthful was usually applied to the plug which had
just been placed. She would press the small ball of
dusty web against the plug, flex her abdomen directly
under her mouth, emit a tiny globule of water apparently
from the anus, and, with the flat-iron movement that
we have elsewhere described for Pompiloides tropicus,
Smooth and rub the material into the plug. This she
did repeatedly. The rubbish she garnered was a motley
mass; it was chiefly dust and spider web, with occa-
Sional small splinters, bits of disintegrated insects and
vegetation, which, combined with the added moisture,
was worked into the nest. Not all of these reinforce-
ments were directed solely to the plug, but many appli-
cations were placed promiscuously about the nest.
To see something entirely new in waspdom, and to
See it done repeatedly, was indeed enough to hold one
alert to catch every detail. To be sure, we had previ-
ously discovered that a near relative, Pompiloides
‘ropicus, likewise bends the abdomen under the body
348 Trans. Acad, Sci. of St. Louis
for the same purpose, and that distant relatives (Ody-
nerus) carry water in their throats. These are near
approaches to the method now discovered, but are not
its counterpart. When I saw that this P. mellipes re-
peatedly left the building by a certain crack, and that
when she returned she was able to gather three pellets
of reinforcing material, I formed the deduction, based
largely on experience with Odynerus, that this was
also a water-carrying species. Yet the puzzling ques-
tion remained unanswered: If she fills her crop with
water, how does it go through the body so quickly to
be emitted from the opposite extremity a few minutes
later? The drop of water was each time handled with
such lightning speed that, I must confess, I might have
been deluded; perhaps the drop came from the mouth,
and the abdomen was so bent as to whip it instantly
from the mouth. I craned my neck all the more; I even
used my magnifying glass, but the work was done too
quickly for the eye to catch the movements.
For an hour while the process of reinforeing contin-
ued, the little mother always went out for water through
the crack nearest the nest and regularly came back
through the same crack. Only once did she enter on
her return through another crack a few inches toward
the north. Did she go from there to her nest? No, she left
mmediately by the way she had éome; presently her
face appeared at the right opening and she proceeded
direct to her nest and resumed her work. Place mem-
ory was again demonstrated.
When at last she seemed to consider her work done
and failed to return, my attention was directed to the
behavior of some half dozen other P. mellipes in the puild-
ing, but of this nest and its contents we shall have more
to say later.
Field Studies of the Non-Social Wasps 349
A second P. mellipes was watched for fifteen minutes
as she nervously walked on the walls or flew from place
to place within the narrow confines of the shack. She
was seen to enter a dozen empty cells of mud wasps,
and one in particular claimed her attention. An hour’s
wait, however, revealed no further activity on the part
of this prospector.
A third individual was seen to return often to a new
and partly wet cell of S. caementarium which was in
course of construction, and I am sure would have started
something had not the owner, returning with her mouth-
ful of mud, routed her with such hearty vigor that P.
mellipes was discreet and never returned. ‘A fourth P.
mellipes on an exploring expedition likewise entered
several old nests but did not remain in any of them.
They generally seem hard to please or for some unex-
Plained reason they do a large amount of prospecting
before they finally select a mud hut to remodel, although
from my standpoint the offerings of available second-
hand domiciles are all about equally good or bad.
A fifth P. mellipes had a sixth one close at its heels
as they both entered through a crack in the wall. The
first one entered an unsealed cell and the other one
still followed; after a few seconds, a wrangling was
heard within and soon after they appeared fighting at
the orifice, the victor threw the rival out, and as she
fell toward the ground she expanded her wings and flew
out at the door. The other remained within. Before
I left an hour later, I inserted a blade of grass in the
cell to ascertain if she was still present, whereupon she
fled in alarm. Late that afternoon I had opportunity
to return, and found her not in, but I did notice that
the opening of the old cell had in the meantime been
enlarged, and the jaw-marks gave evidence that some
350 Trans. Acad. Sct. of St. Louts
of the clay had been bitten away from the periphery.
By the next morning, more of the clay around the open-
ing had been eut away, and inserting a stem of grass
I sensed that some structural work had been done with-
in. Late that afternoon I paused a moment just before
train-time and removed the nest, incomplete as it was,
to take home, and was surprised to see that more clay
had been taken from about the aperture. Looking with-
in, I could see at the far end a completed and open cell.
Now to return to an examination of the first cell,
whose closure I witnessed as recorded above. This
mother had taken the unused cell and modified it in
accordance with the plan illustrated in the diagram
(fig. 39). In cell ‘‘A’’ the inside walls were smooth and
slick, and a black spider of the species Phidippus tri-
punctatus [J. H. Emerton] was within. Like many
other individuals of this species observed, this wasp
had failed to remove all of the legs of its prey, but those
which she had taken off were amputated in a workman-
like manner — that is, with a nice straight cut a few
millimeters from the base. Again, she differed from
others in her work in that the spider’s remaining legs
were not completely paralyzed. How often we read in
the text-books that the wasp stings the prey so that the
body remains fresh food for the young, while the legs
are paralyzed to protect the young from injury. In
this spider the body seemed dead, but the remaining
legs were active and clung to my pencil or finger with
great tenacity. After the lapse of five days it was in
about the same condition of activity as when the nest
was first opened and 15 days later this spider shed its
skin. This little incident may have a deeper meaning,
too. It is possible that this species directs her stings
to affect other parts besides the legs, for what would
Field Studies of the Non-Social Wasps 351
be the utility of stinging so as to paralyze the legs, in
a species which has the habit of cutting off the legs of
its prey? An egg, now discolored, was sealed to the
left lateroventral surface of the abdomen. Cell ‘‘B”’
was likewise smooth on the inside; all of its roughness
faced into the empty cell ‘‘C’’ where it would do no
harm since it was empty. In ‘‘B”’ one spider, a young
Philodromus sp. [J. H. Emerton] was found. This was
dead, bore a dried, shriveled egg upon its dorsum, and
had only three legs. The five that had been removed
were the first, third and fourth on the left side and the
third and fourth on the right. All these seemed to
have been cut at the joint between the coxa and the
femur, leaving clean-cut stumps. ‘‘D’’ was a very large
cell between the last brood cell and the plug, and con-
tained sufficient space for at least three P. mellipes cells,
but without taking advantage of this space she had pro-
ceeded to plug up the orifice. We ought to pause here
to ask how the habit was acquired of sealing up the open-
ing of the tunnel or old cell when her own cells within were
sufficiently sealed, since in many situations of her nest,
as under bark of trees, ete., she cannot seal up the cavity
°r Space containing her cells.
Fig. 39 again shows the great versatility of this little
creature. In this case she did not actually build the
usual thimble-shaped cells, but modified her work to
resemble that of Trypoxylon clavatwm (fig. 54), and
actually built walls to separate the larve instead of
following the more elaborate plan of building separate
cups for each. In contrast with the precaution of leay-
ing an air space, as seen at ‘‘D”’ in fig. 39, and sealing
the opening, we see (fig. 38) where the old mud-dauber’s
cells were completely filled with thimble-like cells of P.
mellipes and in other cases some of them were partly
352 Trans. Acad. Sci. of St. Louis
exposed to view, not having the plug at the opening.
All of this shows that ‘‘habit’’ has not yet bound this
creature to her wheel, and it suggests that this species’
instinct is today in a very unstable condition and will
bear watching for a solution of some of the most per-
plexing problems of psychic evolution.
The promising prospect of wresting from P. mellipes
some of the secrets of her nest-building caused me to
return to Wickes. But on this second visit, in Septem-
ber, the outhouse that had previously been of interest
proved void of P. mellipes. At 5:35, however, almost
twilight, I found in a dark corner of another shed nearby
a P. mellipes atop the nest of a mud-dauber. The nest
was more or less dilapidated, with whole sides torn out
of certain cells. At first this condition aroused little
attention, since I thought the nest had merely met with
some calamity such as an attack by birds, or disintegra-
tion by the elements. Later investigations, however,
showed that this destruction was the work of P. mellipes;
hence I give briefly the condition of the nest when found.
One cell that had been sealed empty had the entire top
removed; a second upper cell had likewise had the top
torn away, but this had been a sealed cell, and the pupal
case of the Sceliphron was hanging over one wall, where
evidently it had been pushed by P. mellipes as she was
demolishing the cell and carrying the mud inside for
her new cells. The two cells underneath were of special
interest. One had the wall about the opening removed,
forming a rather large circular aperture, twice the size
of the original opening. This cell will hereafter be re-
ferred to as ‘‘X.’? The mandible-marks were vividly
etched about the periphery. The other under cell was
the main item of interest, because the P. mellipes was
at the time building her own cells therein of the mud
Field Studies of the Non-Social Wasps 353
which she obtained by breaking down and carrying away
other parts of the old nest. When she had first begun
to make her cells herein, she had evidently gotten her
mud from the very same cell, for enough had been re-
moved about the opening to enlarge it to two or three
times its normal size and the edge was all scarred with
mandible-marks. This cell is known as ‘‘H.’’
P. mellipes was on the nest at the time of its discov-
ery, but left as I approached. After a lapse of nine
minutes she returned, flew into cell ‘‘X,’’ presently
came out and flew into her own cell ‘‘H.’’? She went in
head first; the abdomen partly protruded and showed
every indication that she was at work within the cell.
Evidently when she entered cell ‘‘X’’ she removed a
mouthful of mud from one of the walls, and within the
cell which she had appropriated she was busily fashion-
ing it into her own cells. The next two mouthfuls proved
the correctness of this assertion, for she was next ac-
tually seen biting out mouthfuls of mud from the broken
cell on top and carrying this material into the cell she
was reconstructing. Since these pellets were not car-
ried out, it is evident that they went into her masonry.
Thus she removed six loads from the wall and took
them down into her cell; then she went out and returned
in 12 minutes. Throughout the period of observation
She often left. the nest, undoubtedly for water, and re-
turned each time after an interval of about two min-
utes. The light was dim in the corner of the shed under
the roof where she was at work, and I could not see
whether she moistened the dirt first before removing it
or wet it as she was actually at work biting it out with
her jaws. Of only this much I was certain: that the soil
as she carried it in her mouth was wet and glistening, and
very close scrutiny revealed that the spot where she got
354 Trans. Acad, Sci. of St. Louis
her mouthful of dirt was always dry. Odynerus geminus
was lavish in the use of water, when mouthful by mouth-
ful she bit away the chimney over her nest, but this eco-
nomical little creature either bit away every particle that
contained water or bit out the dry dirt and disgorged
upon it just enough water to make it of the right con-
sistency. She came repeatedly to the wall of the nest ap-
parently empty-handed, made actual mud in her jaws
for short periods and then went away. Working thus,
she continued to remove the foreign portions of the nest,
carry the mud to a certain cell and there to fashion her
own dainty little thimble-shaped cells. Often, as she
worked, her hind legs extended out to view, and fre-
quently I could see her curl the abdomen under the body,
as previously described. Again I tried hard to deter-
mine the question which had puzzled me on previous 0¢c-
casions, whether the water oozes out of the tip of the
abdomen when it is so inflexed or whether the drops come
from the mouth and are whipped into place by the tip
of the abdomen as it begins to rub down the mud, but
again I was unsuccessful in the dim light.
The speed of her work may be judged from the fol-
lowing sample of her comings and goings, and the num-
ber of mouthfuls of mud removed after each trip. She
went out at 5:40 p. m., returned at 5:42; out two min-
utes, then removed 8 mouthfuls of mud and worked it
into her own architecture, consuming 6 minutes at the
task; out again at 5:48, in at 5:49, absent one minute,
removed 7 loads of mud in the same fashion in the fol-
lowing 6 minutes; out for 2 minutes, then transferred
6 loads of mud to her own structure in three minutes;
out again for three minutes, and in the following 5 min-
utes took out 6 loads of dirt.
Field Studies of the Non-Social Wasps 355
The wall-structure of this Sceliphron nest had not
been reinforced in the usual way; consequently it was
thin and quantities of it were required to make the com-
pact cell of P. mellipes. Hence the tops of three large
cells had been demolished and much additional material
taken from the other two cells, to put into the making
of the three stocky little cells which had been built with-
in the one chosen mud-dauber’s cell.
The P. mellipes builder did not each time return direct
to one spot to get her mud, but walked all over the nest
in a searching attitude, sometimes more than once, be-
fore stopping to get her mouthful. Sometimes she went
to the same place on two consecutive trips, but more
often she landed at a new spot each time. ‘At about 6
0’clock the cup-shaped cell was completed very near to
the opening, and the next few loads brought in were used
to fill in the niches about the new cell. When she left
at 6:08 and did not return at the usual time, I suspected,
since her nest seemed completed, that she had gone in
search of a spider. I waited until 7 p. m., when it was
quite dark in the shed, but she did not return. Occa-
sional visits during the two days following showed me
no evidence of the return of the mother. Perhaps she
had met with a tragedy; perhaps I had frightened her
away by trying to observe the progress of her work by
the aid of lighted matehes. The cell which she had
completed before her disappearance was found to be
empty ; the middle one contained a spider, Phidippus sp.
[J. H. Emerton], and had a discolored egg fastened
across the dorsal surface of the abdomen. The remain-
ing cell, the oldest of the three, contained an almost full
8rown P. mellipes larva and a few fragments of spiders.
It was replaced and on September 24 it spun its cocoon.
356 Trans. Acad, Sci. of St. Lows
In the living-room of the club-house this yellow-legged
Pompilid was once seen crossing the floor with her
spider. When I came too near she flew to the window,
but presently she returned slowly and cautiously. Dur-
ing her absence I examined the spider, a young Phidip-
pus [J. H. Emerton] and found that it had only one leg,
a front one; the other seven had been bitten off close
to the body. Thus it seems that each individual P. mel-
lipes trims the legs of her spiders exactly to suit her
own fancy, and not in accordance with any law or fixed
instinct. She left the spider lying on its back; when
she returned she mounted it without changing its posi-
tion, which made the ventral side of the spider rest
against the ventral side of the wasp. Her body, and
that of the spider, are about the same size, so she nicely
_ covered it as she trundled it to a point some three feet
away. Despite the length of her legs she was unable
to lift it from the floor, so it rubbed against the rough
boards until I thought surely it would get a splinter in
its back. She stopped alongside a loose board and with-
out leaving her spider or changing her position in the
slightest degree, she paused and rested for a full half
hour. Sometimes, while still in the same position, she
would lift her hind legs and stroke her body with them
= in toilet-making. The second half hour was drag-
ging wearily on; her capacity for waiting seemed to
exceed mine. I blew my breath gently but directly upon
her; this immediately aroused the antenne to very wild
movements, but soon her agitation cooled and she quieted
them. This, however, did not induce her to move,
but she became quite tame, so that I could approach
her near enough to see that with her mandibles she held
the abdominal tip of the spider as it lay beneath her. Suc-
cessive blowings at length roused her to move on with
Field Studies of the Non-Social Wasps 357
it for a distance of two feet, where she lapsed again into
the same loitering, sitting astride her spider. I wished
to let her alone and observe her natural capacity for
loafing, but I could not remain longer so took both wasp
and spider.
In the foregoing pages we have remarked upon the
unusual versatility of this little wasp. This character-
istic is generally regarded as an advantageous faculty,
but we must not forget that versatility may lead to stu-
pid, labor-wasting ways as well as ingenious, economical
methods. Witness the following story of wasted time
and energy.
This P, mellipes was seen carrying a spider under her
body in the usual way, i. e., venter to venter and the
anal tip grasped in the wasp’s jaws, while the first pair
of legs held it close to the wasp’s body. Thus the wasp
walked actively about on the face of an almost perpen-
dicular bank, and entered about a dozen abandoned
Anthophora burrows, coming out very soon from all ex-
cepting the last; there she remained for some time. I
placed a test-tube over the hole to take her when she
Should emerge, but when she saw the tube she darted
back into the depths of the burrow, but soon again ven-
tured to the orifice. Seeing that something was wrong,
she went back into the hole again, got her spider and
attempted to escape with it, which I permitted. After
that she carried this spider in and out of more than
thirty holes, always holding it in the same manner, going
into one hole twice without seeming to know it, becom-
ing entangled once in a spider’s web, from which she
cleared herself with a few well-placed kicks, once tum-
bling down with her burden from the top of the bank,
but spreading her wings when half way down and re-
gaining her position. Eventually, she came out of one
308 Trans. Acad, Sci. of St. Louis
of the many holes empty-handed. With a probe I found
the hole so shallow that the spider was easily poked out.
It was legless; the legs had all been bitten off very near
to the body. This is my first record of a totally de-
legged spider. The victim, a Pisaurina wndata Htz.
[C. H. Shoemaker], was evidently dead. For the next
half hour she continued to entertain herself by going
into other abandoned holes. It is possible that this par-
ticular wasp had a mud cell hidden in one of the bee-
burrows and, becoming confused at not finding the right
cell, abandoned the prey.
P. mellipes, too, has her enemies. In September, 1919,
a cuckoo-bee, Tetrachrysis pattoni Aaron [S. A. Rohwer],
was reared from one of its cocoons.
A brief summary of the nesting habits of the mem-
bers of this genus, as far as they have been reported,
might be of interest here in a comparative way, although
finer details of the methods of manipulation are needed
before we can get much from a study of their relation-
ships through this behavior.
The pots of Agenia punctum* are made of mud and
are shaped like oval jars, each smaller than a cherry
stone. Those of A. hyalipennis affect a conoid form,
narrow at the base and wider at the mouth like a primi-
tive drinking cup. The nests of both species are glazed
on the inner surface; this makes them waterproof.
(Fabre, The Mason Wasp, p. 84, and Sharp, Insects, Pt.
2, pp. 105-106.)
Agenia carbonaria contrives a nest much like a wide-
mouth bottle. It appears that this insect has not learned
*Many of these insects were formerly known under the generic
name of Agenia, but since the name was preoccupied in the Hemiptera,
he genus is now named Pseudagenia (Banks, Journ. N. Y. Ent. Soe
19:221, 1911). No attempt is made in these reviews to standardize the
nomenclature, but each name is the one used by the author cited.
Field Studies of the Non-Social Wasps 359
the secret of kneading its material with saliva, so its pots
have not the proper permanency, and for this reason
they are not placed in exposed situations. (Step, Mar-
vels of Insect Life, p. 426; Sharp, Insects, Pt. 2, pp.
105-106.)
Pseudagenia adjuncta makes tiny mud cells in the old
cocoons left in the nests of Trypoxylon politum, and it
also plasters larger mud cells to the same nests. (Rau,
Journ. Anim. Behay., 6; 42, fig. 21, 23.)
Agenia subcorticalis builds nests in old Pelopoeus
cells, taking the dirt from the walls of the nests in which
She is building, moistening the dirt with water as she
works. (Hartman, Bull. Univ. Texas, 6 : 51.)
Pseudagenia blanda builds stout cells of mud, cylin-
drical and thick walls, ranged alongside each other,
Sometimes as many as four cells in a nest; they are
placed in the fissure of a vine or at the base of a tree.
(Williams, loc. cit., p. 97.)
Pseudagenia makilingi makes a two-celled mud nest
which is hidden in a curled-up leaf. (Williams, loc. cit.,
p. 97.)
Agenia bombycina makes clusters of little mud cells
% crevices. (Peckham, Wasps Social and Solitary, p.
4.)
Pseudagenia architecta makes twin cells of mud. (Rau,
Wasp Studies Afield, p. 84.) These nests are probably
smilar to those described by the Peckhams (Wise. Geol.
& Nat. Hist. Surv., 2; 165-166), but they do not men-
tion any attachment of the cells in pairs. Ashmead
(Psyche 7; 66. 1896) records that the nests are thimble-
Shaped, and are found under bark, logs and rocks. This
brief description by Ashmead is applied also to three
other species, A. bombycina, A. corticalis and A. mellipes.
360 Trans. Acad, Sci. of St. Louis
Agenia variegata. This species is recorded as bur-
rowing in the ground. (Riley, Am. Nat. 8; 8. 1874.)
This, however, needs verification, since no other record
shows similar behavior for members of this genus;
Bouvier says (Psychic Life of Insects, p. 190) that A.
variegata closes the entrance of her nest with balls of
spider web which she compresses into a tight wad, al-
though he does not state what kind of nest this species
makes.
Pseudagenia nyemitawa. This wasp plasters a two- or
three-celled nest on trunks or logs. (Williams, loc. cit.,
p. 98.)
Pseudagenia caerulescens makes a little mud nest
within the silken retreat of a jumping spider, the web
of which is usually in a crack in a bamboo stump. (Wil-
liams, loc. cit., p. 100.)
Pseudagenia sp. had enlarged a burrow in a honey-
combed log. Within, a short tunnel was found which
contained three cells, each supplied with a spider. (Wil-
liams, loc. cit., p. 101.)
Pseudagenia macromeroides shows indications of a
semi-social habit; the mud cells are placed in groups
in sheltered places. (Williams, loc. cit., p. 102.)
Pseudagenia okowa is a twig-nesting wasp. She par-
titions the hollows of slender twigs into 2, 3 or 4 cells,
and stores them with the delegged spiders. The parti-
tions are of mud, and in addition the outer one is smeared
with a gummy substance (Williams, loc. cit., p. 103).
Thus we see that the genus Pseudagenia, more often
called Agenia, although it belongs to a family of wasps
that dig burrows, comprises for the most part potter
wasps that build neat pots of various shapes a¢
_ cording to the species, in various localities, with or with-
Field Studies of the Non-Social Wasps 361
out garnishments, according to whether the nest is in
the open or in protected situations. All of these species
carry mud from afar to build their nests. Sometimes,
however, some individuals of P. mellipes eliminate this
work by building their nests in a daub of dry mud and
carrying only the water to this point of operations, thus
Saving a great amount of the work. Not all individ-
uals of P. mellipes behave in this manner; this variation
shows at least that this habit is new to the species and
probably in an incipient stage of higher development.
One must not overlook, too, that Hartman has discov-
ered that Agenia subcorticalis uses the cells of Pelopoeus
merely as cavities in which to build her own small cells
of the ancestral type. Thus she may have as many as
five of her own cells inside a single chamber of a mud-
dauber’s nest. Indeed, A. subcorticalis too goes a step
farther and not only closes each of her individual cells,
but builds a plug over the opening to the large chamber,
thus offering to her enemies an additional rampart.
The dirt is taken from the very nest in which she is
building her own; she gnaws off pellets, after having
moistened the dirt with water from her gullet.
When one sees the variations in nesting habits within
each of these two species, A. Mellipes and A. subcorti-
calis, and compares the behavior of the various species
within the genus, one cannot help but look toward the
day when naturalists will be able to show the graduated
Series from the nest of most primitive structure to one
of great complexity and the relationship between the
nesting habits and anatomical characteristics.
362 Trans. Acad, Sci. of St. Louis
CHAPTER III.
Tue Grass-Carrier Wasp, Chlorion (Isodontia)
auripes Fern.
The behavior of this very interesting creature was
incompletely told in ‘‘Wasp Studies Afield,’’ pp. 203-
206, and Trans. Acad. Sci. 25; 199-201, 1926. Since that
time I have been able to add, piecemeal, several details
of its life history. In 1917 we occasionally encountered
the striking spectacle of a wasp flying high with a long
blade of grass or strand of excelsior in its jaws. We
finally traced these wasps to carpenter-bee burrows in
wooden beams, where they were carrying in this mate-
rial, and so ended our information on the habits of the
species. ‘‘Thus the season ended without our having
ascertained whether they used this material for bedding,
for food, or as a plug to close the orifice.’’ With our
curiosity thus aroused, we were ever on the watch for
their burrows in the wood.
We have previously quoted Packard, who tells how
these wasps use analagous material for plugging their
nests. More recently we were pleasantly surprised to
find a note by F. X. Williams* which has some bearing
on the phylogeny of this habit. He records that a slen-
der wasp from Philippine Islands (species not recorded)
“‘of the Zsodontia group, provided with slim mandibles
and legs unfitted for digging, was observed gathering
tomentum or wool-like material from the under side of
a green leaf, and she evidently makes use of some pre-
existing hollow as a nesting place, dividing the cells
therein with the material gathered.’’
My first interesting discovery was a pine board in
horizontal position, above a doorway in an abandoned
*Philippine Wasp Studies, p. 119, 1919.
Field Studies of the Non-Social Wasps 363
cabin near Kirkwood. A typical carpenter-bees’s old
hole had been filled in with grass, tightly packed. The
diameter of the hole at the opening was 7/16 inch (see
1 in fig. 41). This opening led to a gallery going east
for a few inches (see 2 in fig. 41), then turned back
sharply and went toward the west (just below 3) for 8
inches.