Title: Florida Entomologist
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098813/00132
 Material Information
Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1974
Copyright Date: 1917
Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
 Record Information
Bibliographic ID: UF00098813
Volume ID: VID00132
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: Open Access

Full Text



Volume 57, No. 3 September, 1974


TURNIP-.I FD. S.-.'SliMp/inM .'U the'fin In ,'It./ h I\'n it,,, D .''L .VI',' -S p. au l
G ,. .ind ('"/,th .ll /th.ul, 217
BI.TI.FR. (; I. AND F. L. WATSON-.4 Technlqqt for DeptLrniningn Ith linrl ,,
Ihti /,pnifnit ,f L guus he.-perus in Flhctclitin, Tmniperatii re.- 225
JOHN-SON. (C -Tu.Ti oninic Kel-, and Dist.rihutiinol Pattern-, for VNea'r,. h Sp.i ,r.
,f'Caloptervx Dar-ell'/f'l. 2.31
WADDILL, V., AND M. SHEPAHD--Bt.,/Iog) '1 a Predlnt r.., Stinikbug, Sturetrus an-
chorago. J-Heiniptera Pentaeiriinel 249
MO'CKFORD. E L.-Th_' Echmepter'yx hageni ('oimple~ (PPsrcoptera Lrpidtp.uit-.
drelJ in Florida 255
LEVY'. R.. H. A. VNRINSVELT. AND H. L. CROIMROv-Reoatue ('Cirenrantion re,
alaiot and Tiocta Eh.ementt,- in adultt ttand Intnulatur Stage. of the Re./ Im.
portedt Fire- .A4 rilrninted h\ I[,in Indrwicd X. -Ra\ F/t-,,rescencnr 269
REI.NERT. J. A -TropHti S.,,1 l'i.hra ri tndl Sulhtrn C/rinc/h Bug Control in
Florida 275
Kl-H. L. P. C. E. ALLEN.. J. W\ KINMh RIU.H. AND L. . KIITERT-.4 S tnri.v
Fungi .4 -wcitated ./ith the Lr, eip.i P. Plecia nearctica. in F/rl,,id 281
W\(LFENBAR(,ER. D. A.. (. CNTi. A.A. GCiERR4., .1. G. POMNONl1. S. H ROBINSON,
.AND R. D. GARCIA-(-Chem)ri,-tr/cint Al.4titly rA .t Co (mpMtpOund, A i.4gn.t the
TOhat co Bui (rnim 287
( RAHAMN. AND J R. RAULSIt. N-The Thnic.i Ruui,.rm on St. <'r'ii.\. 1'. .S.
Iirgin I*/.tnd.r : Hr.os Pltint.l. P't.lnatiuoi S aurt tind Eltitunati 297
LFV\ R.. H. L. CINIMR)V. ND IJ A. CORNEl I -ma,,t unl Traot Elemenbn a. Bi-,
predictlors oeqf R dio tinhluol -In .,/ced In,., i. r .s i,';//i .3 I.J
LF.C.AIo. G. L.-Ine rrete in Peoptdort.i ,,i Crn\" ptni.ites pu-illus ond C. turcicus
1n Diel of, .\V'itural Pro, unt. ,309
GRISSFLL. E. E.-.4A I t Dihrmah\-, ih h a i i. ti'. .\'re eif Spec,.'.s (He
ot'ii npteru* Piet ,nntil/t. Pi f .3 13
Y' I N,. D. G.. AND C H P.- F'TERh'-LutzoImyi~ cirrita n p /r,..i* (',,i,,j thl it ith t
\'r, S.'t ,n m it, i te (;,1.11, ti ( i r r-.i '. h i,.it,, F'/ii h f ,,it,, nune I .321
AI.I.F.N. G .ANwl A Si.\ .E ..i-GLi[.iiii--- ,.,,r, .. ,,/ Mn.i ,..p,rni/d, in Solenopis
richteri and Solenop.i-. ,i p m 'rinmn\ Wi a,.4 r32i ii,, 327
PORTER, C. C.-A .\'i Trath\ sphyru.. thc. Phlnrc. I (r'tup r',m ri Fl. rtidia Hy-
menoptera: I: hIn, unit.',ll/t.i 331
Notes andAbstracts .............. 230, 267, 268, 274, 280, 285, 286, 296, 302, 308, 325, 326, 330
Obituary: Kenneth E. Bragdon ................. ...... .............. .......... ... .... .. ...... 224

Published by The Florida Entomological Society


President .......... ......... .................... ......... .................. ............ W G G enung
Vice-President ...................................... ........... R. M. Baranowski
Secretary ........ ........ ........ ................... ........ ..... ............. .. F. W M ead
Treasurer .......................................... ......................... D. E. Short

C. S. Lofgren
Other Members of Executive Committee.......... W.. de
A. B. Selhime
H. D. Bowman
J. R. Strayer


Editor ...................................... .. .. ... .... ... .......... .. S. H K err
Associate Editors ............................................................. .............. R. E. W oodruff
J. E. Lloyd
H. V. Weems, Jr.
Business M manager ... ............ ............. ................. ....... D. E. Short

THE FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and
December. Subscription price to non-members $7.50 per year in advance; $2.00 per copy.
Entered as second class matter at the post office at Gainesville, Florida.
Manuscripts and other editorial matter should be sent to the Editor, Entomology
Department, University of Florida, Gainesville. Subscriptions and orders for back
numbers are handled by the Business Manager, Box 12425, University Station, Gaines-
ville, Florida 32601. The Secretary can be reached at the same address.
When.preparing manuscripts, authors should consult "Instructions to Authors", Vol.
56, p. 364 (1973), and examine recent issues for details of form and style.
The page charge is $5.00 per page, partial pages proportionally. One page of tables is
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This issue mailed November 15, 1974

The Florida Entomologist



Department of Entomology, Clemson University,
Blackville, S. C. 29817


Variations of D-Vac, sweep, and ground cloth methods were compared as
sampling tools for estimating density of populations of insects in soybeans.
Insects sampled included the Mexican bean beetle, Epilachna varivestis
Mulsant; the green cloverworm, Plathypena scabra (F.); the soybean looper,
Pseudoplusia includes (Walker); the corn earworm, Heliothis zea (Boddie);
the velvetbean caterpillar, Anticarsia gemmatalis Hubner; the bean leaf
beetle, Cerotoma trifurcata (Forster); and certain geocorids and nabids.
Ground cloth methods were the most efficient based on numbers collected,
coefficients of variability, and time requirements. Sweeping across and
through 2 rows was adequate for most species and more efficient than sweep-
ing along 1 row. Variations of the D-Vac method were generally ineffective for
the insects sampled in this study.

Published comparisons of sampling techniques for obtaining estimates of
numbers of insects in soybeans are limited. Soybean insects in Minnesota were
sampled by fumigation-cage and sweeping techniques, and species which were
most abundant were underestimated by sweeping (Kretzschmar 1948). Boyer
and Dumas (1963) described a method whereby insects were shaken from
plants onto a cloth spread on the ground between rows. They indicated that
this method was satisfactory for the various species, except for adults of the
three-cornered alfalfa hopper, Spissistilus festinus (Say). Boyer (1967) later
suggested sweeping for adults of S. festinus and plant examination for
Pedigo et al. (1972) compared fumigation-cage, sweep-net, pitfall trap,
shaking over a ground cloth, and D-Vac as sampling procedures for green
cloverworm, Plathypena scabra (F.), in Iowa. The fumigation-cage was the
most precise technique but sweep-net gave greater relative net precision
(based on precision and cost). Pitfall traps and D-Vac samples were the least
precise for this species. The shake sample technique, utilizing a ground cloth,
was not studied sufficiently for proper evaluation.
In general, insect numbers on soybeans have been reported as numbers per
foot of row by dislodging insects and counting from a ground cloth or as
numbers per sweep(s). The vacuum-net method has often been used for
sampling parasites, predators, and small phytophagous species.
In 1968 and 1969 we made limited comparisons of the efficiency of D-Vac,
sweep, and ground cloth methods for sampling certain phytophagous species.

'Technical contribution No. 1112, Department of Entomology and Economic Zoology, South
Carolina Agricultural Experiment Station, Clemson. Published by permission of the Director.
Received for publication 9 January 1974.

Vol. 57, No. 3, 1974

218 The Florida Entomologist Vol. 57, No. 3, 1974

The study was expanded in 1972 to include comparisons of 2 variations of each
of these methods. No attempt was made to determine absolute density of the
species (or stages) sampled. Insects sampled were the Mexican bean beetle,
Epilachna varivestis Mulsant; the green cloverworm; the soybean looper,
Pseudoplusia includes Walker; the corn earworm, Heliothis zea (Boddie);
the velvetbean caterpillar, Anticarsia gemmatalis Hubner; the bean leaf
beetle, Cerotoma trifurcata (Forster); and certain geocorids and nabids.

Vacuum-net samples were taken with a D-VacR model 24 machine with a
13 in. diam collection head. For the vertical variation of the D-Vac method
(DVV), the collection head was held vertically and placed down over plants as
the operator walked along the row; whereas with the horizontal variation
(DVH), the collection head was held horizontally within the row. Sweep
samples were taken with a heavy 15 in. diam sweep net. For the across
variation (SAC), the sampler swept across and through 2 rows as he walked
between them; whereas with the along variation (SAL), sweeps were taken
along (on alternate sides of) 1 row while walking beside it. In 1968 and 1969,
live insects were counted directly from nets in the field for D-Vac and sweep
methods. In 1972 insects were killed by enclosing net contents in killing jars
with ethyl acetate. Samples were then emptied into polyethylene bags and
stored in a freezer for later processing. Processing was accomplished by (1)
drying samples under a lamp in the laboratory, (2) separating insects from
trash with a camel's hair brush, and (3) counting insects with the aid of a
dissecting microscope.
Ground cloth samples were collected on and counted directly from a 4 ft x
40 in. piece of heavy white cloth in the field. For the beat variation (GCB), the
cloth was placed between two 40 in. rows with a person at each end. The
foliage of 1 row was carefully moved aside away from the cloth and foliage of
the adjacent row was bent over the cloth and beaten vigorously 6 to 8 times by
a downward motion with the hands and forearms. After quickly pushing
the 4 ft section of row aside, insects were counted on the cloth. A 4 ft
sample was taken in 1969. In 1968 and 1972 the process was repeated in
each plot for an 8 ft sample. For the shake variation (GCS), the cloth was
placed between 2 rows both of which were shaken vigorously together over
the cloth by striking 6 to 8 times with the hands and forearms. Both rows
were quickly pushed aside and insects dislodged from the two 4 ft sections
were counted on the cloth. In both ground cloth methods care was taken
to count quick-moving insects first. To avoid recounting and to prevent
escape, geocorids and nabids were crushed on the cloth as they were
Row width in all fields was 40 in. All sampling methods were replicated 4
times in a randomized block design. Data were transformed prior to analysis
using \x + Variety and growing conditions of the soybeans in sampled
fields are shown in Table 1.


Data comparing D-Vac horizontal (DVH), sweep across (SAC) and ground
cloth beat (GCB) in 1968 and 1969 for sampling Mexican bean beetle, green
cloverworm, soybean looper, corn earworm, bean leaf beetle and velvetbean

Turnipseed: Sampling Soybean Insects



Year no. Variety Growing conditions

1968 1 Hardee /2 ft high; standing well; early bloom
2 Hardee 2/2-3 ft high; standing well; mid pod set
1969 1 Hardee 21/2-3 ft high; standing well; early bloom
2 Hardee 2/2-3 ft high; standing well; early bloom
1972 1 Bragg 3-31/ ft high; standing well; late bloom
2 Hardee 3-31/2 ft high; standing well; mid pod set
3 JEW 46 22-3 ft high; standing well; pods 60% filled

caterpillar are shown in Tables 2 and 3. GCB of 8 ft of row produced greater
mean numbers of all species than SAC 15 times. In 1968 GCB 8 ft and SAC 15
times yielded similar numbers of adult Mexican bean beetles. Although only 4
ft were sampled by GCB in 1969 (Table 3), results were similar. DVH of 20 ft of
row was third in numbers of each species (or stage) collected.


MEAN NUMBERS OF E. varivestis, P. scabra, P. includes AND

Avg no. insects per sample**

Sampling Field 1 (16 Aug) Field 2 (6 Sept)
and size* E. varivestis

Adults Larvae P. scabra P. scabra P. includes H. zea

DVH 20 ft 4.5a 9.3a 6.5a 6.3a 0.5a 3.8a
SAC 15 times 16.0b 49.8b 8.5a 19.5b 2.8b 20.5b
GCB 8 ft 16.8b 128.5c 13.5a 28.8b 7.5c 39.5c

*DVH 20 ft=D-Vac horizontal of 20 row ft; SAC 15 times=sweep across 15 times; GCB 8
ft = ground cloth beat of 4 row ft.
**Means followed by the same letter are not significantly different at the 5% level (Duncan's
multiple range test).

In the expanded study of 1972 (Table 4), field populations of insects men-
tioned earlier, except bean leaf beetle, were sampled by the 6 previously
described methods. The ground cloth methods (GCB and GCS) usually gave


The Florida Entomologist

similar results and yielded greater mean numbers with smaller coefficients of
variability than the other sampling methods. However, GCB yielded sig-
nificantly greater mean numbers of soybean loopers (Fields 1 and 2), green
cloverworms (Field 1) and nabids (Field 3) than GCS. Fifteen sweeps by either
SAC or SAL generally followed ground cloth methods in numbers collected
and, with the exceptions of geocorids and cloverworms, SAC collected more
insects than SAL. The D-Vac methods (DVV or DVH, Fields 1 and 2)
consistently produced lowest mean numbers with highest variability.

TABLE 3.-MEAN NUMBERS OF E. varivestis, P. includes, C. trifurcata

Av. no. insects per sample**
Field 1 Field 2
Sampling E. varivestis -A
method vests P. P. C. A.
and size* Adults Larvae Pupae incl. incl. trif. gem.
DVH 20 ft 19.5a 4.3a 0.8a 0.3a 1.0a 3.5a 0.8a
SAC 15 times 110.Oc 19.5ab 2.3a 2.8b 2.8ab 11.5b 4.0ab
GCB 4 ft 54.5b 31.3b 10.3b 7.0c 5.3b 24.5c 6.3b

DVH 20 ft = D-Vac horizontal of 20 row ft; SAC 15 times = sweep
across 15 times; GCB 4 ft = ground cloth beat of 4 row ft.
** Means followed by the same letter are not significantly different at the
5% level (Duncan's multiple range test).

Selection of a sampling method for soybean insects would depend upon the
species present in the complex and upon the particular species for which
population estimates are desired. Additionally, the size or developmental
stage of a species may dictate selection of a sampling method. If, for example,
numerous pupae of the Mexican bean beetle were present in a field, then GCB
would probably produce a more accurate measurement of numbers than SAC
or DVH (Table 3). Also, efficiency of the method must be considered; with
efficiency taking into account numbers collected, variability, and time or
expense involved.
For the insects included in this study, the ground cloth methods (sampling
8 ft of row) consistently yielded greater mean numbers and generally had
lower coefficients of variability than either variation of sweeping 15 times or
vacuuming 20 or 25 ft. GCB collected significantly higher mean numbers of
the soybean looper, the green cloverworm and nabids than did GCS. Beats
may have produced higher mean numbers of these insects than shakes
because: (1) greater numbers of loopers, which hold rather tenaciously onto
foliage, were dislodged by the vigorous beating of plants downward compared
to shaking in a less vertical motion; (2) shaking resulted in more lateral
dislodging than beating downward causing some insects to fall on the soil

Vol. 57, No. 3, 1974

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Turnipseed: Sampling Soybean Insects

away from the ground cloth; and (3) more fast-crawling cloverworms or
quick-moving nabids may have escaped detection when specimens from two 4
ft sections of row were counted at a time (GCS) rather than one 4 ft section at
a time (GCB).
Sweeping across was generally an effective method, but processing of
samples in the laboratory was time consuming. If a few target species were
counted directly from the net, efficiency of this method would be greatly
The D-Vac methods were not efficient tools for sampling the insects
studied. However, the vacuum-net is quite effective for sampling small,
quick-moving insects such as leafhoppers, whiteflies or hymenopterous
parasites (Turnipseed, unpublished data).


Boyer, W. P. 1967. Survey method for three-cornered alfalfa hopper (Spissis-
tilus festinus) in soybeans in Arkansas. USDA Coop. Econ. Insect Rep.
Boyer, W. P., and W. A. Dumas. 1963. Soybean insect survey as used in
Arkansas. USDA Coop. Econ. Insect Rep. 13:91-92.
Kretzschmar, G. P. 1948. Soybean insects in Minnesota with special reference
to sampling techniques. J. Econ. Ent. 41:586-91.
Pedigo, L. P., G. L. Lentz, J. D. Stone, and D. F. Cox. 1972. Green cloverworm
populations in Iowa soybean with special reference to sampling
procedure. J. Econ. Ent. 65:414-21.


224 The Florida Entomologist Vol. 57, No. 3, 1974

Kenneth Edward Bragdon, Special Inspector (retired), Division of Plant
Industry, Florida Department of Agriculture and Consumer Services (for-
merly the State Plant Board), was born on 10 December 1886 in Sullivan,
Maine. He died 23 February 1974 in Savannah, Georgia. He was employed as
an agent by the State Plant Board on June 1915 at Gainesville, Florida. Prior
to his employment with the State Plant Board in 1915, he held the position as
teacher-superintendent of schools and as a clerk with the Puerto Rican
government. He was promoted to general grove supervisor 16 March 1916 and
discovered 2 of the several infestations of citrus canker. In the spring of 1918 he
was appointed as an agent with the Bureau of Entomology, U. S. Department
of Agriculture for the sweetpotato weevil program. On 30 November 1919 he
was appointed county agent for Brevard County. Before returning to the
State Plant Board 1 July 1941 to work with the white-fringed beetle project in
Florala, Alabama, he held the position of production manager with the Winter
Haven Citrus Growers Association. He was then transferred to Winter Haven
in the Grove Department on 26 June 1944 and was appointed district inspector
at Vero Beach 1 January 1946. On 15 January 1953 he became assistant to the
Grove Inspector and on 1 July 1953 was appointed Special Inspector. He
retired 1 January 1956.
Kenneth Bragdon was recognized for his early work on the citrus canker
and sweetpotato weevil and later the whitefringed beetle and tristeza
programs. He was the last surviving charter member of the Florida En-
tomological Society, the second secretary-treasurer for the Society, and the
second business manager for The Florida Entomologist, then known as The
Florida Buggist. He was the 26th president of the Society in 1942. The Society
recognized Kenneth Bragdon for having served faithfully for over 30 years by
presenting him with the Honors Award and conferring him with honorary
membership in 1954. He was a Mason for over 50 years and was awarded the
50-year membership citation.
A Masonic graveside service was held 1 March 1974 at Lakeside Memorial
Park Cemetery, Winter Haven. He is survived by 2 sons, Kenneth Paul Brag-
don, Savannah, Ga.; Dennis E. Bragdon, Winter Haven, Fla.; and a daughter,
Juanita Baker (Mrs. Alfred), Winter Haven, Fla.
H. A. Denmark
Division of Plant Industry
Fla. Dept. Agric. & Consumer Serv.

S. *

The Florida Entomologist



A computer program was prepared that utilizes developmental data de-
termined in the laboratory at constant temperature to determine the duration
of the stages of Lygus hesperus Knight at fluctuating temperatures. This
program provides the basis for a generalized insect development and popula-
tion model (WATBUG).

Laboratory studies of the effect of constant temperatures on the develop-
ment of eggs and nymphs of Lygus hesperus Knight showed an increase in the
rate of development with an increase in temperature (Butler and Wardecker
1971). Analyses of population trends of L. hesperus in California alfalfa fields
indicated that heat input and temperature extremes played a dominant role in
determining the rate of population increase (Butler 1971). Stitt (1940) made
observations of the incubation period of the egg of L. hesperus and the dura-
tion of the instars in an outdoor insectary but these results are not amenable
for the prediction of development under other outdoor conditions because the
mean daily temperature sometimes does not take into account the fluctua-
tions that occur in hourly temperatures. If we are to utilize field studies under
fluctuating temperatures or predict future field populations, a technique is
needed to convert developmental information obtained in the laboratory at
constant temperatures to the developmental rate that prevails at fluctuating
temperatures. The present paper reports such a technique and demonstrates
that the observed duration of the different stages of L. hesperus at fluctuating
temperatures compared favorably with those predicted by a computer
program based on rates determined at constant temperatures.

The concept basic to the present model is that the rate of development
varies with temperature, any delay in the change when temperature changes is
assumed to be negligible. The proportion of development of a stage of an insect
during a given time period can be determined from the reciprocal of the time of
development determined at constant temperatures (Fye et al. 1969). In our
technique a computer program, written in FORTRAN, uses tables of
development rate versus temperature to calculate the duration of the
different stages of an insect under almost any given temperature flux. A flow
graph showing the logic of the program is given in Fig. 1.

SThis work was supported in part by grants from the Cooperative State Research Service and
Cotton Inc. In cooperation with the Arizona Agr. Exp. Sta. Received for publication 20 Sept. 1973.
SEntomology Research Division, Agr. Res. Serv., USDA, Western Cotton Research Laboratory,
Phoenix, Ariz. 85040.
:1 Formerly with the University of Arizona Agricultural Experiment Station, Tucson, Ariz. 85721.

Vol. 57, No. 3, 1974


226 The Florida Entomologist Vol. 57, No. 3, 1974

Initialize the day and read tables of development vs.

Increment the day by 1
Read the temperature for the day sampled at even times
Initialize to 1st time period of the day

Increment period of day by 1
Initialize stage of insect being considered

Increment stage of insect life
Calculate average development for period of time
Multiply by period
Initialize to 1st group of this stage
Add to column of development accumulation
Is stage of development complete? ----yes

no initialize development and
__change to next stage
no Is this all groups of this stage?
o yes
o-- Is this all of stages of insect life?
o yes
no Is this the last period of day?
I yes
Insert any new stages
no I
nIs this the last day of simulation?
I yes


Fig. 1.-Flow graph of logic of WATBUG program.

One set of data used to test this technique was obtained when 3,150 eggs
laid on 33 days and the nymphs from these eggs were reared in an outdoor
insectary at fluctuating temperatures during a 71-day period from late
January through April 1967 (Butler and Wardecker 1971). Another set of data
was obtained from observations of nymphs hatched in the laboratory, dusted
with colored pigment, and transferred to cages constructed of 1-qt cartons
that were placed over alfalfa plants growing in the field. The cages were
brought to the laboratory daily and the nymphs examined for pigment to

__ __ __

__ _ __ __

Butler: Modeling Rate of Development



15 20 25 30 35

Fig. 2.-Comparison of the observed and calculated
stage of Lygus hesperus in an insectary.

duration of the egg

determine when molting occurred. Nymphs that had not molted were re-
turned to the field. Thermograph records were obtained by placing a ther-
mograph probe in one of the cages. Overheating in the direct sun was
prevented by conducting the test in a large screen cage with a shade cloth
across the top. Daily temperatures fluctuated from 17 to 300C, and one daily
high reached 350C.


The duration of the egg stage in an outdoor insectary varied from 14.2 to
30.2 days. The program was set to calculate the duration of the egg stage by
using the observed 3-hr temperatures. The calculated values had a range of 14
to 31 days. Fig. 2 shows the relation between a regression line of the observed
and calculated observations and one of equal values.
The duration of the 5 nymphal stages in the outdoor insectary was also
determined and compared with the duration calculated with the program
utilizing observed 3-hr temperatures. Fig. 3 shows the relationship between
the observed and the calculated times. The program was most successful in
estimating the duration of the-lst, 3rd, 4th, and 5th stages, but most calcula-
tions were within the mean + 1 day, the increment of time at which observa-
tions of change of instar were observed.
The duration of the nymphal stages in the field in cages on alfalfa during
the heat of the summer was determined and compared with the results ob-
tained with the program (Table 1). Most calculated values were within the


w 25-


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C) 15-
0 J

228 The Florida Entomologist Vol. 57, No. 3, 1974


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0 9 10

6 0 0 7,


Fig. 3.-Comparison of the observed and calculated duration of the
nymphal stages of Lygus hesperus in an insectary.

mean + standard deviation of the observed values. However, in this test, the
limited range of temperatures and the small number of observations was not
sufficient to justify the calculation of regression equations.


The duration of the life stages of insects in an insectary or in the field is
difficult to predict by using an average mean temperature that prevailed
during the stage because' this average temperature does not take into account
the fluctuations in the daily temperatures that may occur, particularly during
the winter. However, the technique discussed here allows determination of the
duration of the egg and nymphal stages of L. hesperus for almost any com-
bination of fluctuating temperatures. This technique was further developed
into a population model (WATBUG) by F. L. Watson (1973, unpublished
dissertation) and used to predict an optimal control law for L. hesperus on

Butler: Modeling Rate of Development



No. Observed mean Calculated
Date nymphs no. days +.SD no. of days

1st stage

June 21
July 1

June 11
July 8

June 13

June 11

June 13

2nd stage

3rd stage

4th stage

5th stage


Butler, G. D., Jr. 1971. Fluctuations of populations of Lygus hesperus Knight
in California alfalfa fields. Pan-Pac. Ent. 47:123-6.


The Florida Entomologist

Butler, G. D., Jr., and A. L. Wardecker. 1971. Temperature and the develop-
ment of eggs and nymphs of Lygus hesperus. Ann. Ent. Soc. Amer.
Fye, R. E., R. Pantana, and W. C. McAda. 1969. Developmental periods for
boll weevils reared at several constant and fluctuating temperatures. J.
Econ. Ent. 62:1402-5.
Stitt, L. L. 1940. Three species of the genus Lygus and their relation to alfalfa
seed production in southern Arizona and California. USDA Tech. Bull.
741, 19 p.

SOCIAL HYMENOPTERA-(Notice.) When disturbed mechanically,
minor workers of Camponotus (Colobopsis) saundersi and of C. sp. near
saundersi contract their gaster until it bursts at an intersegmental fold. The
mandibular glands, which extend throughout the whole body, also burst
releasing large quantities of a whitish yellow (C. saundersi) or bright yellow
(C' sp. near saundersi) secretion. The secreted fluid is very sticky and at-
tacking ants are unable to move when contaminated with it. The term "au-
tothysis" (greek.: self sacrifice) is proposed for the phenomenon. Oecologia
(Berl.), 1974, 14, 289-294; U. and E. Maschwitz, Univ. Frankfurt/M.


Vol. 57, No. 3, 1974

The Florida Entomologist



Department of Zoology,
University of Florida, Gainesville

This paper gives taxonomic keys for adults of the 5 nearctic species of
Calopteryx damselflies and completes a 3-title series on distributions of
calopterygid species occurring in the United States. Separate male and female
keys provide species determinations. The accompanying text clarifies in-
traspecific variability, female morphs, areas of sympatry for similar spe-
cies, and reviews the taxonomic uncertainty of C. amata and C. angusti-
pennis. Distributions, totally within the U. S. and Canada, appear for
each species by appropriate state or province subunits, and focuses atten-
tion on related ecological questions. Flight season data reveal little or no
seasonal differences between species. A review of early southwestern rec-
ords for C. maculata appears, and existing literature notes provide a tenta-
tive identification of the early type-locality of C. angustipennis represent-
ing the single species report for Georgia.

This paper gives a key for adult determinations, distributional patterns
and flight seasons for the Nearctic species of Calopteryx damselflies. The
Calopterygidae, represented in the United States and Canada by only 2
genera, Hetaerina and Calopteryx, constitute our larger and more colorful
damselflies. Behaviorists and ecologists have increasingly adjusted their
studies to the readily-observed calopterygid behavior and such efforts will
doubtless improve our knowledge of evolution and speciation in these dam-
selflies. An objective examination of species distributions revealing likely.
allopatries and sympatries is timely, and such data appeared for Hetaerina
and, in part, for C. dimidiata in earlier reports (Johnson 1973a,b).
Current literature recognizes 5 nearctic species of Calopteryx, maculata
(Beauvois) 1805; dimidiata Burmeister 1839; aequabilis Say 1840; angus-
tipennis (Selys) 1853; amata Hagen 1890. These large, colorful insects at-
tracted collectors early in the entomological explorations of North America,
and, as the above dates show, only 1 recognized species description appeared
after 1853. Hagen's synopsis of nearctic Calopteryx, dated 1889 but officially
published on 4 January 1890 (F. M. Carpenter, personal communication 1973),
identified and allocated synonyms and amplified species descriptions. Hagen's
paper clearly indicated his appreciation of a species' potential for geographic
variability. He reduced C. hudsonica, a name proposed earlier by him in 1875,
to a subspecies of C. aequabilis and relegated C. apicalis to a synonym of C.

SResearch Associate, Florida State Collection of Arthropods, Bureau of Entomology, Florida
Department of Agriculture and Consumer Services, Gainesville 32601.

Vol. 57, No. 3, 1974

The Florida Entomologist

dimidiata. Both decisions arose from a consideration of geographic variation.
Larger collections over the respective geographic areas proved his interpreta-
tion of C. apicalis valid 83 years later (Johnson 1973b). Hagen's same paper
included the description of C. amata based on four males and females respec-
tively from Dublin, New Hampshire. He had reservations about C. amata due
to his limited material of C. angustipennis available for comparison. When
Hagen, in Cambridge, Massachusetts, wrote the 1890 synopsis, the only known
male of C. angustipennis (the type, presumably from Georgia) was in the
British Museum and he possessed only 2 C. angustipennis females. The
description of C. angustipennis appeared as Sylphis angustipennis Selys 1853
for the female. Hagen recognized the association of male and female in his 1861
"Synopsis of North American Neuroptera" and commented more fully on the
decision in the 1890 Synopsis of North American Calopteryx concluding, "The
material known for C. angustipennis is decidedly not adequate; if a larger
number should prove the difference given for C. amata not persistent, the two
species will belong together." Needham and Heywood (1929), 39 years later,
referred to C. amata as . doubtfully distinct."; however Walker's (1953)
study, the major diagnostic work following Needham and Heywood, covered
northern colonies recognized as C. amata. Actually, the name C. amata
appears more frequently in the literature, regional lists, etc. than C. angus-
tipennis, a species still uncommon today in most collections. The only authors
directly addressing variability in nearctic Calopteryx following Hagen were
Cockerell (1913) and Kennedy (1917a, 1918) on subspecies of C. aequabilis,
Huggins (1927) on a subspecies of C. maculata, and Johnson (1973b) on
seasonal and geographic variation in C. dimidiata. Needham and Heywood
(1929) provide the only key, long out-of-print, for the 5 nearctic species. Its
brief style omits variation and recognition of female morphs, thus allowing
erroneous determinations, and the short distributions accompanying species
accounts require considerable updating.
Mate recognition by Calopteryx involves color patterns and flight behavior
(Bucholtz 1955; Pajunen 1966). Species criteria consist, therefore, of wing
patterns, relative wing width, color of the males' ventral abdominal segments,
occasionally stigma size in females and body lengths. Conventional structure
of male abdominal appendages, penis, and female ovipositor, often of value in
Odonata, have little taxonomic distinction in the nearctic species. Color pat-
terns frequently require a maturation period for full expression and, in con-
junction with body size, may vary both seasonally and geographically. Female
morphs involve one form typically mimicking the male (the andromorph or
homeomorph) and a second morph (the heteromorph) differing distinctly. At
least in C. dimidiata, females may be andromorphic for only the hind wing
pair (Johnson 1973b). These attributes become apparent only in large samples
and complicate the writing of taxonomic keys.
Walker (1953), Johnson (1972) and most entomological guides provide
definitions and illustrations of key characters. Notes on variability leading to
possible confusion in determinations follow the keys.

la. Wings dark brown, often appearing opaque black (except
occasionally isolated cell(s); wing length equals approx-
imately 3X the greatest width ............ .............. ....... ...... maculata

Vol. 57, No. 3, 1974


Johnson: Keys and Distribution of Calopteryx

b. Wings clear or slightly overcast with amber or brown; a
dark brownish to black band may occur on the apical
end of the wing extending a variable distance toward the
nodus; wing length greater than 3.5X the greatest width ................ 2
2a. All wings with a dark brown band in the apical end; wing
length = 3.5 to 4X the greatest width ............. .................... ............... 3
b. All wings clear of brown to black pigment or only the hind
wings with an apical dark band (although often a very
pale brown); wing length greater than 4X greatest width................... 4
3a. Apical dark band (usually brown) of hind wing distinctly
wider (measured parallel to wing margin) than fore wing
band; sternum of abdominal segment 10 white or pale cream
co lo red ...... ............ .... ....... .... ...... .. ........ .. ................ a eq u a b ilis
b. Apical dark bands of fore and hind wings approximately
equal in length, rarely varying over 2 mm; sternum of ab-
dominal segment 10 dark green or black ................................. dimidiata
4a. Hind wings only with an apical brown band, apparently
never approaching the near-black color seen in some speci-
mens of above species, often very pale; sternum of abdominal
segment 10 with irregular whitish spot; ventral surface of
metathorax pale, becoming pruinose in older specimens ............. amata
b. All wings clear (rarely with slight amber overcast); sternum
of abdominal segment 10 and ventral surface of metathorax
variable, see following notes .......................................... angustipennis

la. Wing length approximately 3X the greatest width; wings
light to dark brown over entire surface (except occasional
and irregular cells), sometimes darker apically .................... maculata
b. Wing length greater than 3.5X the greatest width; wings
clear, occasionally with amber or brown overcast, or clear
basally and with a dark brown apical band ...................... ............. 2
2a. Wing length 3.5 to 4X the greatest width; hind margins of
wings rounded and not parallel to front margins; wings
often with apical dark brown bands; pre-antennal ridge
with some dark green or black color; abdomen usually less
th an 40 m m .......................................... ...... ............ ............ ............... 3
b. Wing length greater than 4X the greatest width; hind mar-
gins of wings rather straight, nearly parallel to the front
margin in the apical half; wings without apical dark brown
bands or very weakly expressed in the hind wings only; pre-
antennal ridge with pale coloration; abdomen usually greater
th an 40 m m ....... ....... ... .......... ...................... ............................. 4
3a. Labrum and labium largely pale; pre-antennal ridge with
some pale markings, often entirely pale; apical brown bands,
if present in wings, usually wider in the hind wings (meas-
ured parallel to wing margin) than fore wing bands.............. aequibilis
b. Labrum, labium and pre-antennal ridge predominantly dark
in color; if apical brown bands present in wings, they are

The Florida Entomologist

approximately equal in length, rarely varying more than
2 m m ................ ...... ................... ........................ dim idiata
4a. Stigma present but may be crossed by veins and sometimes
very small; wings overcast with amber, occasional specimen
with pale apical hind wing band similar to male ........................ amata
b. Stigma absent; wings clear or very slightly touched with
amber, no apical band in any known specimen .............. angustipennis

The wide-ranging Calopteryx maculata stands distinctly apart in color
and morphology from its congeners, and distributional patterns supplement
the key in recognizing the other species. Coexistence of the superficially
similar C. dimidiata and C. aequabilis exists in only a small northeastern
region (Figure 2), and only C. amata of the C. amata-C. angustipennis pair
ranges north of Pennsylvania (Figure 3). Southward, both C. amata and C.
angustipennis occur, though uncommonly, and our limited knowledge of
southern colonies leaves some uncertainty of the species criteria.
Large samples of C. angustipennis and C. amata over their geographic
range are still unavailable; however, criteria used for these species remain
diagnostically recognizable in the available material (16 males and 6 females
of C. angustipennis from Kentucky, North Carolina, Pennsylvania, South
Carolina, Tennessee, and West Virginia; 15 males and 30 females of C. amata
from Massachusetts, New Hampshire, New York, North Carolina, Pennsyl-
vania, Tennessee, and West Virginia). Determinations of older specimens in
the series were by E. B. Williamson, E. M. Davis, and P. P. Calvert. Carl Cook,
Leonora K. Gloyd, Paul Harwood, M. J. Westfall, the author and others have
determined more recent materials. While the sample is not large by statistical
standards, the above odonatologists concur in determinations, and the sample
represents the best comparative series available. In this material, all male C.
amata possess the apical brown band of the hind wing. The band is present but
pale in young specimens and intensifies somewhat with advancing age. The
silhouetted photograph in Robert (1963), Fig. 41, p. 67, of a male C. amata
wing from Quebec, is essentially devoid of the hind wing band; nonetheless,
Robert's key utilizes the band as a character. Walker's (1953) photograph,
plate 8, clearly shows the band in a Quebec male; Robert's specimen was
perhaps very young or the photograph lost contrast in reproduction. The dark
band appears in only 1 C. amata female of the sample, is very pale, and the
specimen is mature. The only other nearctic Calopteryx species known to
possess apical dark bands on only the hind wings is a female morph of C.
dimidiata (Johnson 1973b); however, a similar study of C. aequabilis may
reveal similar females. The wing. band of C. amata is pale in some mature
specimens, and, judging from behavioral work on European and Eurasian
species by Buchholtz (1955), it is doubtful that such bands function well in
species recognition.
The most definitive attributes for C. angustipennis are the key characters
given above; however, a puzzling variation exists in the series for this species.
The white or pale cream-colored sternum of abdominal segment 10 is a con-
stant male attribute of species possessing it (e.g., all examined males of C.
maculata, C. aequabilis and C. amata have the whitish sternum and all C.
dimidiata males have a greenish-black sternum). Reproductive behavior in-
volves display of the white spot, when present, to the female (Williamson 1904;

Vol. 57, No. 3, 1974

Johnson: Keys and Distribution of Calopteryx

Johnson 1962; Pajunen 1966). In C. angustipennis from Kentucky, Pennsyl-
vania, and Tennessee, the sternum of abdominal segment 10 is black, and 4
additional males in the British Museum from Pennsylvania also possess black
sterna (P. Ward, personal communication 1973). Specimens from North
Carolina, South Carolina and West Virginia have a whitish sternum of ab-
dominal segment 10. Unfortunately, the terminal segments are missing on the
male holotype (from Georgia) in the British Museum (Kimmins 1969). Inten-
sity of black pigmentation increases with age; however, these differences exist
in clearly mature specimens. Since the abdominal white spot is a known
component of behavioral releasers in species-specific behavior, it is natural to
question its variability in C. angustipennis. Specimens are somewhat more
numerous than during Hagen's day; however, questions now include behavior.
Relative to this trait, one may repeat Hagen's observation, the knowledge of
C. angustipennis is "... .decidedly not adequate".

The complete range of Western Hemisphere Calopteryx species occurs in
the United States and Canada with 2 species, C. angustipennis and C.
dimidiata, confined to the eastern United States. Calopterygid damselflies are
stream to riverine species and will typically occur only about such habitats
within the specific ranges given below. Distributions appear by county or
parish for each state; the Canadian distributions follow Walker (1953) giving
counties for all provinces except Manitoba and Saskatchewan where the few
sites have other locations. These localities form the range maps in Fig. 1-3
where each point lies approximately in the center of the represented county.
Johnson (1973b) reported a detailed distribution for Calopteryx dimidiata
earlier and only the subsequent new record follows. Literature listed
chronologically, personal communication sources, and collections) providing
documentation for the data follow county lists. I cite only papers giving
specific localities. Initial reports of a species for some states do not therefore
appear. Collections cited carry the following abbreviations: CJ Coll.-author's
coll., GHB Coll.-G. H. Bick's Coll., PM Coll.-Paul Milliotis' Coll., Clem. U.
Coll.-Clemson University Coll., Corn. U. Coll.-Cornell University Coll., U. C.
Coll.-University of Connecticut Coll., U. A. Coll.-University of Arkansas
Coll., U. N. Coll.-University of Nebraska Coll., U. M. Coll.-University of
Michigan Coll., U. V. Coll.-University of Vermont Coll., INHS Coll.-Illinois
Natural History Survey Coll., and FSCA-Florida State Collection of

Calopteryx aequabilis distributional records.
UNITED STATES: California: Humboldt County. Kennedy (1917).
Colorado: Boulder and Larimer counties. Cockerell (1913); Kennedy (1918).
Connecticut: Litchfield, New Haven, New London and Tolland counties.
Woodruff (1914); Garman (1927); U. C. Coll. Idaho: Benewah, Blaine, Latah,
Owyhee, Twin Falls and Valley counties. Barr (personal communication
1973); FSCA. Illinois: Boone and Kane counties. INHS Coll.; U.M. Coll.
Indiana: Elkhart, Fulton, Starke and White counties. Montgomery (1941).
Iowa: Black Hawk County. Wells (1917). Maine: Androscoggin, Kennebec,
Knox, Lincoln, Oxford, Penobscot and Washington counties. Howe (1917);
Borror (1951). Massachusetts: Berkshire, Essex, Franklin, Middlesex, Norfolk
and Worchester counties. Howe (1917, 1919, 1921); PM. Michigan: Alger,

o 0.
** o*f .
-V^*.*,. *& */^ W _

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1**.** .
** *4 *r n *I,. *
j. i^* *^"^*..X \' ..

. : ,.*? ^9 .* 7
.*.*:.:.; * : ;
- *** : C * ** ** __i.* .. *-
* '1 ^ (.* ,,* **. *. Y ^ *
I ,-- I* *I, *** ..
0 o

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*0 0 *, *.

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The Florida Entomologist


Fig. 3. Distributional patterns of Calopteryx amata and C. angustipennis.

Alpena, Arenac, Baraga, Berrien, Cheboygan, Clare, Crawford, Delta,
Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Gratiot, Houghton,
Huron, Ingham, Iron, Isabella, Jackson, Kalamazoo, Keweenaw, Lake, Liv-
ingston, Marquette, Midland, Missaukee, Oakland, Ontonagan, Oscoda, Ot-
sego, Presque Isle, Roscommon, Schoolcraft, Shiawassee, Washtenaw and
Wexford counties. Kormondy (1958). Minnesota: Aitkin, Anoka, Carlton,


Vol. 57, No. 3, 1974

Johnson: Keys and Distribution of Calopteryx

Cass, Clay, Clearwater, Douglas, Hennepin, Hubbard, Lake, Lyon, Martin,
Mille Lacs, Morrison, Nicollet, Norman, Olmstead, Pine, Ramsey, Red Lake,
Redwood, Scott, St. Louis, Stearns, Wabasha and Winona counties. Wilson
(1909); Clausen and Hamrum (personal communications 1973); FSCA. Mon-
tana: Flathead County. Newell (1970). Nebraska: Logan and Sioux counties.
FSCA; U. N. Coll. Nevada: Elko County. La Rivers (1940). New Hampshire:
Belknap, Carroll, Grafton, Hillsboro, Merrimack, Rockingham and Strafford
counties. Howe (1917); White and Morse (1973); PM. New Jersey: Passaic
County. Howe (1921). New York: Bronx, Clinton, Essex, Herkimer, Oswego,
Suffolk and Tompkins counties. Calvert (1895a); Needham (1903, 1928);
FSCA. North Dakota: Grand Forks and Pembina counties. Post (personal
communication 1973); FSCA. Ohio: Portage and Summit counties. Borror
(1937). Oregon: Benton, Deschutes and Malheur counties. Kennedy (1915,
1918); FSCA. Pennsylvania: Bucks, Centre, Mifflin and Union counties.
Beatty, Beatty and Shiffer (1969, 1970). South Dakota: Bennet and Meade
counties. Balsbaugh (personal communication 1973); Corn. U. Utah: Box
Elder and Weber counties. Brown (1934); Waage (personal communication
1974). Vermont: Caledonia, Chittenden and Orleans counties. Howe (1917); U.
V. Coll.; FSCA. Washington: King and Yakima counties. Kennedy (1915,
1918). Wisconsin: Dane, Milwaukee, Saint Croix, Sheboygan, Vilas and
Washington counties. Muttkowski (1908, 1909); Howe (1921). Wyoming: Te-
ton County. Bick (1972).
CANADA: Manitoba: Treesbank and Winnipeg. Walker (1953). New
Brunswick: Carlton, Charlotte, Kings, Sunbury, Westmoreland and York
counties. Walker (1953). Northwestern Territories, District of Mackenzie,
Fort Simpson. Waage (personal communication 1974). Northern-most
Calopteryx record in the western hemisphere and not shown on distribution
map. Nova Scotia: Annapolis, Halifax, Hants, Kings and Pictou counties.
Walker (1953). Ontario: Brant, Carlton, Cochrane, Dufferin, Frontenac, Grey,
Hastings, Huron, Lanark, Leed, Middlesex, Nipissing, Oxford, Parry Sound,
Peel, Prince Edward, Renfrew, Simcoe, Timiskaming, Waterloo, Wellington
and York counties. Walker (1953); Waage (personal communication 1974).
Quebec: Brome, Gatineau, Labelle, Liette and Vercheres counties. Walker
(1953); Waage (personal communication 1974). Saskatchewan: Waskesiu
Lake. Walker (1953).
Calopteryx amata distributional records.
UNITED STATES: Connecticut: Litchfield County. Woodruff (1914). Maine:
Oxford and Somerset counties. Harwood (1959); FSCA. Maryland: Garrett
County. Howe (1921). Massachusetts: Middlesex County. Howe (1917); PM
Coll. New Hampshire: Carroll, Cheshire, Coos, Grafton and Merrimack
counties. White and Morse (1973); PM Coll. New York: Essex, Saint
Lawrence and Sullivan counties. Needham (1928). North Carolina: Bun-
combe, Henderson, Macon and Transylvania counties. FSCA. Pennsylvania:
Cameron, Centre, Clearfield, Fayette, Forest, Huntingdon, Luzerne, Potter,
Sullivan, Susquehanna, Tioga, Union, Warren and Westmoreland counties.
Ahrens, Beatty, and Beatty (1968); Beatty, Beatty and Shiffer (1969). Ten-
nessee: Blount County. Goodwin (1968). West Virginia: Grant, Pocahontas,
Raleigh, Randolph and Tucker counties. Cruden (1962); Harwood (personal
communication 1973); FSCA.
CANADA: New Brunswick: Sunbury and York counties. Walker (1953).
Quebec: Brome County. Walker (1953).

Calopteryx angustipennis distributional records.
UNITED STATES: Georgia: Burke-Screven counties. (?) See Discussion.
Indiana: Crawford County. Williamson (1917). Kentucky: Breckinridge, Ed-

The Florida Entomologist

monson, Grayson, Hart, Marion and Rockcastle counties. Resner (1970);
Cook (personal communication 1973); FSCA. North Carolina: Haywood
County. CJ. Ohio: Ashland and Richland counties. Williamson (1899); Borror
(1937). Pennsylvania: Butler, Cameron, Centre, Fayette, Franklin, Somerset,
Union and Westmoreland counties. Williamson (1899); Ahrens, Beatty and
Beatty (1968); Beatty, Beatty and Shiffer (1969). South Carolina: Oconee
County. Clem. U. Coll. Tennessee: Fentress and Sevier counties. Trogdon
(1961); FSCA. West Virginia: Grant, Hampshire, Mineral, Nickolas, Pendle-
ton and Webster counties. Harwood (1972); FSCA.

Calopteryx dimidiata distribution supplement.
Tennessee: Pickett County. CJ.

Calopteryx maculata distributional records.
UNITED STATES: Alabama: Baldwin, Covington, Lee and Shelby counties.
Huggins (1927); Wright (1943). Arkansas: Johnson, Marion, Montgomery,
Phillips, Stone, Washington and Yell counties. Adams (1900); U. A. Connect-
icut: Fairfield, Litchfield, New Haven, New London, Tolland and Windham
counties. Howe (1917); Garman (1927); U. C. Delaware: Kent, New Castle and
Sussex counties. Roback and Westfall (1967); Kelsey (personal communica-
tion 1973). Florida: Alachua, Baker, Bradford, Calhoun, Clay, Columbia,
Escambia, Gadsden, Hamilton, Hardee, Highlands, Hillsborough, Jackson,
Jefferson, Lake, Leon, Levy, Liberty, Madison, Marion, Okaloosa, Orange,
Osceola, Polk, Putnam, Santa Rosa, Seminole, Suwannee, Taylor, Union,
Volusia, Wakulla, Walton and Washington counties. FSCA; CJ. Georgia:
Clarke, Dooly, Jefferson, Lee, Rabun and Wayne counties. Root (1924); Byers
(1931); FSCA. Illinois: Carroll, Champaign, Christian, Clark, Coles, Cook, De
Kalb, Hardin, Iroquois, Mason, McHenry, McLean, Ogle, Peoria, Piatt, Pike,
Pope, Stephenson, Union, Vermillion, Will and Winnebago counties. Garman
(1917); FSCA; INHS Coll. Indiana: Allen, Benton, Boone, Carroll, Clark,
Clinton, Crawford, Elkhart, Fountain, Fulton, Gibson, Hamilton, Harrison,
Howard, Jackson, Jasper, Johnson, Kosciusko, La Porte, Madison, Marshall,
Monroe, Montgomery, Noble, Orange, Parke, Porter, Posey, Pulaski, Ran-
dolph, Starke, Tippecanoe, Union, Vigo, Warren, White and Whitley counties.
Williamson (1900); Montgomery (1935, 1937, 1941, 1951, 1953, 1955, 1971);
FSCA. Iowa: Black Hawk, Boone, Muscatine, Polk, Story and Tampa coun-
ties. Elrod (1898); Wilson (1912); Wells (1917). Kansas: Douglas, Ellis, Kiowa,
Phillips, Rawlins, Riley, Rooks, Rush, Shawnee and Summer counties. Ken-
nedy (1917); Roback and Westfall (1967); FSCA. Kentucky: Adair, Allen,
Ballard, Barren, Bell, Breckinridge, Bullitt, Butler, Carter, Casey, Cum-
berland, Edmonson, Fayette, Fulton, Graves, Green, Greenup, Hart, Hender-
son, Jefferson, Kenton, Knott, Lewis, Logan, Lyon, Marion, McCreary,
Meade, Metcalfe, Monroe, Muhlenberg, Nelson, Ohio, Oldham, Pike, Pow'ell,
Pulaski, Rockcastle, Taylor, Todd, Trigg, Union, Warren, Wayne and Whitley
counties. Resner (1970); Cook (personal communication 1973); FSCA.
Louisiana: Beauregard, Bossier, Claiborne, East Baton Rouge, East
Feliciana, Jackson, La Salle, Lincoln, Natchitoches, Quachita, Rapides, St.
Tammany and Washington parishes. Bick (1957). Maine: Androscoggin,
Franklin, Hancock, Kennebec, Knox, Lincoln, Oxford, Penobscot, Waldo and
Washington counties. Borror (1944, 1951); Milliotis (personal communication
1973). Maryland: Anne Arundel, Baltimore, Charles, Frederick, Montgomery
and Prince Georges counties. Root (1923); Huggins (1927); Donnelly (1961);
FSCA. Massachusetts: Barnstable, Bristol, Essex, Franklin, Hampshire,
Middlesex, Norfolk, Plymouth, Suffolk and Worchester counties. Howe (1917,
1919, 1920); Gibb and Gibb (1954); PM Coll.; FSCA. Michigan: Alcona, Alger,
Allegan, Alpena, Arenac, Barry, Benzie, Berrien, Branch, Cheboygan, Clare,

Vol. 57, No. 3, 1974

Johnson: Keys and Distribution of Calopteryx

Crawford, Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Gratiot,
Houghton, Huron, Ingham, lonia, losco, Isabella, Jackson, Kalamazoo,
Kalkaska, Kent, Keweenaw, Lake, Lapeer, Lenawee, Livingston, Mackinac,
Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Mis-
saukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland,
Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Presque Isle, Ros-
common, Saint Clair, Saint Joseph, Schoolcraft, Shiawassee, Tusocla, Van
Buren, Washtenaw, Wayne and Wexford counties. Kormondy (1958).
Minnesota: Anoka, Cook, Fairbault, Goodhue, Hennepin, Lake, Lyon,
Nicollet, Olmstead, Pine, Ramsey, Rock, Scott, Steele, Wabasha, Washington
and Winona counties. Wilson (1909); Whedon (1914); Clausen and Hamrum
(personal communications 1973). Mississippi: Claiborne, Covington, Forrest,
George, Hancock, Harrison, Jackson, Lafayette, Lamar, Marion, Marshall,
Noxubee, Pearl River, Perry, Stone and Tishomingo counties. GHB.; FSCA.
Missouri: Barry, Boone, Carter, Greene, Jasper, Johnson, Pulaski, Shannon,
Stoddard and Warren counties. Williamson (1932); Roback and Westfall
(1967); CJ. Montana: Flathead County. Newell (1970). Nebraska: Antelope,
Cass, Cuming, Douglas, Lancaster and Lincoln counties. U. N. Coll. New
Hampshire: Carroll, Cheshire, Coos, Grafton, Hillsboro, Merrimack,
Rockingham and Strafford counties. Howe (1917); White and Morse (1973).
New Jersey: Camden, Cape May, Mercer and Ocean counties. Roback and
Westfall (1967); FSCA. New York: Clinton, Erie, Essex, Franklin, Herkimer,
Madison, Oneida, Schenectady, Suffolk, Tompkins, Ulster and Wyoming
counties. Needham (1928); FSCA. North Carolina: Ashe, Cherokee, Durham,
Graham, Henderson, Macon, McDowell, Mecklenburg, Orange, Swain, Tran-
sylvania, Wake, Wilkes and Wilson counties. Brimley (1903); Byers (1931);
FSCA. Ohio: Adams, Allen, Ashland, Ashtabula, Auglaize, Brown, Butler,
Carroll, Champaign, Clark, Clinton, Coshocton, Cuyahoga, Darke, Delaware,
Erie, Fairfield, Franklin, Gallia, Geauga, Greene, Guernsey, Hamilton,
Highland, Hocking, Holmes, Huron, Jackson, Lake, Licking, Logan, Lorain,
Lucas, Marion, Montgomery, Morrow, Paulding, Perry, Portage, Ross,
Trumbull, Union, Vinton, Warren, Wayne and Williams counties. Borror
(1937, 1938, 1942); Montgomery (1943); Harwood (1960); Roback and Westfall
(1967). Oklahoma: Adair, Alfalfa, Caddo, Choctaw, Cleveland, Comanche,
Craig, Delaware, Ellis, Harper, Kay, Marshall, McCurtain, Murray, Nowata,
Oklahoma, Osage, Ottawa, Pawnee, Payne, Pittsburg, Pushmataha,
Sequoyah, Washita, Woods and Woodward counties. Bick and Bick (1957).
Pennsylvania: Allegheny, Armstrong, Beaver, Berks, Blair, Bucks, Butler,
Centre, Chester, Clinton, Crawford, Cumberland, Delaware, Elk, Erie,
Fayette, Forest, Franklin, Fulton, Greene, Huntingdon, Indiana, Lancaster,
Lawrence, Lebanon, McKean, Mifflin, Monroe, Montgomery, Northampton,
Perry, Philadelphia, Pike, Potter, Snyder, Somerset, Sullivan, Susquehanna,
Tioga, Union, Venango, Warren, Washington, Wayne and Westmoreland
counties. Ahrens, Beatty and Beatty (1968); Beatty, Beatty and Shiffer (1969,
1970). Rhode Island: Kent County. Howe (1917). South Carolina: Allendale,
Charleston, Florence, Greenville, Greenwood, Horry, Laurens, Lexington,
Newberry, Oconee, Orangeburg and Sumter counties. Montgomery (1940);
Roback and Westfall (1967); Clem. U.; FSCA. South Dakota: Bennet County.
Balsbaugh (personal communication 1973). Tennessee: Anderson, Blount,
Campbell, Cheatham, Cocke, Coffee, Cumberland, Davidson, Dickson, Fen-
tress, Greene, Hardin, Hawkins, Jackson, Johnson, Marion, Montoe, Mont-
gomery, Obion, Overton, Pickett, Roane, Sevier, Sullivan, Trousdale,
Washington and Williamson counties. Williamson (1923); Trogdon (1961).
Texas: Anderson, Angelina, Bastrop, Bowie, Cherokee, Collin, Dallas, Den-
ton, Grayson, Gregg, Grimes, Hemphill, Houston, Marion, Montgomery,
Nacogdoches, Robertson, Rusk, Shelby, Walker and Wood counties. Johnson
(1972). Vermont: Bennington, Caledonia, Chittenden, Grand Isle, Orleans,
Rutland and Washington counties. Howe (1917); U. V. Coll.; PM Coll. Vir-

The Florida Entomologist

ginia: Fairfax, Montgomery, Roanoke and Washington counties. Williamson
(1903); Gloyd (1951); Donnelly (1961); Johnson (1962). Washington, D. C.:
Hagen (1874); Donnelly (1961). West Virginia: Boone, Braxton, Brooke,
Cabell, Doddridge, Gilmer, Grant, Greenbrier, Hampshire, Hardy, Jefferson,
Kanawha, Lewis, Lincoln, Logan, McDowell, Marion, Marshall, Mason,
Mercer, Mineral, Monongalia, Monroe, Ohio, Pendleton, Pleasants,
Pocahontas, Preston, Putnam, Raleigh, Randolph, Ritchie, Roane, Summers,
Taylor, Tyler, Upshur, Wayne, Webster, Wetzel, Wood and Wyoming coun-
ties. Huggins (1927); Cruden (1962); Harwood (personal communication
1973); FSCA. Wisconsin: Burnett, Door, Forest, Langlade, Marinette, Mil-
waukee, Oconto, Sheboygan, Vilas and Waukesha counties. Muttkowski
(1908); Clausen (personal communication 1973); FSCA.
CANADA: Manitoba: Waugh. Walker (1953). New Brunswick: Carleton,
Charlotte, Northumberland, Queens, Sunbury and Westmoreland counties.
Walker (1953). Nova Scotia: Annapolis, Digby, Halifax and Pictou counties.
Walker (1953). Ontario: Algoma, Muskoka, Nipissing, Parry Sound and Wa-
terloo counties. Walker (1953). Quebec: Gatineau, Huntingdon and Labelle
counties. Walker (1953).
Seasonal flight times clearly do not isolate the species where their ranges
overlap as particularly noted by Woodruff (1914) and Wells (1917) for
northern areas. I have observed C. maculata and C. dimidiata co-existing in
Florida, on occasion taking both species in one swing of the net. Walker (1953)
gave 27 May to 10 September for C. maculata in Canada, and it may occur as
adults the full year in south-central Florida during mild winters. Johnson
(1973b) found the season for C. dimidiata in New Jersey (near its northern
boundary) as 28 May to 25 August, and it approaches a full-year existence in
Florida (26 February to 5 November). Walker's (1953) dates for C. aequabilis
in Canada are 3 June to 2 September. An earlier date, 6 May (New Jersey)
exists in the FSCA series; however, no later specimens occur than reported for
Canada. The known seasonal ranges for C. amata and C. angustipennis are 31
May (Massachusetts) to 6 August (Tennessee), and 18 April (Georgia-Ab-
bott's type) to 5 July (North Carolina) respectively. Harwood's experience in
West Virginia (personal communication 1973) suggested local populations of
C. amata and C. angustipennis have short flight seasons.


Colonies of C. maculata have occurred over most riverine conditions in the
eastern half of the United States and southeastern Canada (Fig. 1). Pollution
has doubtlessly reduced populations, and the species fails naturally to
colonize more open, non-forested streams of the west. Nonetheless, 3 reports
of C. maculata appeared for California, Nevada and New Mexico, and have
been cited by later authors. The earliest report was by Calvert (1895b) for
California giving only, "One male, California . ." The specimen was in a
consignment sent to Calvert for study from the California Academy .of Science
Collection. Fire later destroyed the collection and Calvert's specimen will
remain a mystery. Muttkowski (1910) first utilized Calvert's record, and
Kennedy (1917b), who had considerable prior field experience in California,
first questioned its validity. Seeman (1927) and Smith and Pritchard (1956)
nevertheless included the species in the California fauna based apparently on
Calvert's early report. R. Garrison (personal communication 1973) searched
collections of the University of California without finding California
specimens and no such specimens exist to my knowledge. Smith and Pritchard

Vol. 57, No. 3, 1974

Johnson: Keys and Distribution of Calopteryx

also give Nevada for C. maculata, once again without specific localities, and
substantiating specimens are unknown. I. La Rivers (personal communication
1973) never encountered the species in his experiences with Nevada Odonata.
Without additional data I assume the California and Nevada reports reflect
errors. The New Mexico report originated with Kennedy (1917b) who gave a
general distribution statement for species occurring in Kansas, and it formed
the basis of Montgomery's (1947) reference of C. maculata from New Mexico
30 years later. I collected Odonata regularly for 5 years in New Mexico
without finding C. maculata and can find no evidence of New Mexico
specimens. Kennedy was at the University of Kansas when writing his 1917
Kansas paper and George Byers (personal communication 1973) stated that a
specimen of that period, with a handwritten label, still exists in the Snow
Museum (University of Kansas) Collection. The locality is "Wolfeboro, N.H."
but is "readable" as "N.M.". I suggest Kennedy misread the label. The western
Montana record by Newell (1970) is based on a badly broken specimen sent to
the FSCA. This Montana colony would appear to be distinctly isolated from
the major gene pool of the species and warrants study for possible divergence.

The range of C. aequabilis forms roughly a horizontal band over the
northern United States and southern parts of central and eastern Canada
(Fig. 2). Factor(s) determining its southern boundary pose an interesting
ecological question, and Martin's (1939) life cycle studies suggest possible lines
of investigation. The spotty distribution in the west appears to involve
isolated populations, and the resulting subspecific names (C. a. californicum
Kennedy 1917; C. a. coloradicum Cockerell 1913; C. a. yakima Hagen 1890; C.
a. hudsonica Hagen 1875) reflect variation largely in the pattern of wing
bands. Western samples are too small to evaluate quantitatively. Recent
Colorado specimens are unknown since Cockerell's work.

The distributional hiatus in the range of C. dimidiata mentioned by
Johnson (1973b) for North Carolina and Virginia narrows specifically to Vir-
ginia by incorporating data in the addenda of that paper. The new Tennessee
record places C. dimidiata quite close to Kentucky, Burmeister's 1839 type
locality, and the reservations on that locality raised in the above paper have
less justification.
The ranges of C. amata and C. angustipennis shown in Fig. 3 correlate
largely with mountain or up-land type streams. Records of C. angustipennis
to the west in Indiana and Ohio represent old, single event occurrences, and
south of West Virginia, the data show a spotty distribution of low-density
colonies. The single Georgia locality for C. angustipennis comes from the 1853
type. The larger Odonata collections possess no additional Georgia specimens,
and I have sought possible specimens from local insect collections in Georgia,
Alabama and South Carolina to no avail. Efforts to specifically trace the type
locality follow.
Selys (1853) gave only "Georgia" for distribution with the original
description and P. Ward (personal communication 1973) informed me that the
type label gave only Georgia. Hagen (1861) identified the collector as Abbott,
and, in 1863, sketched John Abbott's contributions to entomology. From
Abbott's notes for C. angustipennis, he gave "April 18, Briar Creek, and rarer".
Presumably, Hagen studied Abbott's notes when he visited London in 1857.
Since the type was still the only male known to Hagen in 1890, there is no


The Florida Entomologist

question that the data represent other specimens. Murphy (1945) gave a brief
history of Abbott's contributions to ornithology indicating that he spent most
of his life in Screven County, Georgia, in the late 1700's. The major drainage
creek in Screven County appears on maps currently as Brier Creek, also
occurring largely in Burke County. I suggest, therefore, the type locality is the
Burke-Screven County area. The region is currently unlike the known habitat
of C. angustipennis but its range is possibly diminishing, judging from the
single early collections noted in Indiana and Ohio to the west. Abbott ap-
parently found it uncommon ("rarer") on Brier or Briar Creek in the 1700's.

The following individuals provided unpublished localities for the
Calopteryx species or otherwise answered specific questions; these contribu-
tions significantly increased the data and clarified otherwise puzzling points:
P. J. Clausen, Univ. of Minnesota; R. L. Post, North Dakota State Univ.; I. La
Rivers, Univ. of Nevada; W. F. Barr, Univ. of Idaho; E. U. Balsbaugh, Jr.,
South Dakota State Univ.; L. P. Kelsey, Univ. of Delaware; C. L. Hamrum,
Gustavus Adolphus College; R. Garrison, Univ. of California; G. Byers, Univ.
of Kansas; L. K. Gloyd, Univ. of Michigan; M. J. Westfall, Jr., Univ. of
Florida; B. E. Montgomery, West Lafayette, Indiana; F. M. Carpenter, Har-
vard Univ.; P. Ward, Br. Mus. Nat. Hist.; P. Corbet, Univ. of Waterloo; J.
Waage, Brown Univ.; P. Harwood, Ashland, Ohio; C. Cook, Center, Ken-
tucky; P. Miliotis, Dunstable, Mass. Paul Laessle drafted the figures.


Adams, C. C. 1900. Odonata from Arkansas. Ent. News 11:621-622.
Ahrens, C., G. H. and A. F. Beatty. 1968. A survey of the Odonata of western
Pennsylvania. Proc. Pa. Acad. Sci. 42:103-109.
Beatty, G. H., A. F. Beatty, and C. N. Shiffer. 1969. A survey of the Odonata of
central Pennsylvania. Proc. Pa. Acad. Sci. 43:127-136.
Beatty, G. H., A. F. Beatty, and C. N. Shiffer. 1970. A survey of the Odonata of
Eastern Pennsylvania. Proc. Pa. Acad. Sci. 44:141-152.
Bick, G. H. 1957. The Odonata of Louisiana. Tulane Stud. Zool. 5:71-135.
Bick, G. H., and J. C. Bick. 1957. The Odonata of Oklahoma. Southwestern
Nat. 2:1-18.
Bick, G. H., and L. E. Hornuff. 1972. Odonata collected in Wyoming, South
Dakota, and Nebraska. Proc. Ent. Soc. Wash. 74:1-8.
Borror, D. J. 1937. An annotated list of the dragonflies (Odonata) of Ohio.
Ohio J. Sci. 37:185-196.
Borror, D. J. 1938. Additions to the Ohio list of dragonflies (Odonata). Ohio J.
Sci. 38:307-310.
Borror, D. J. 1942. New Records of Ohio dragonflies (Odonata). Ohio J. Sci.
Borror, D. J. 1944. An annotated list of the Odonata of Maine. Can. Ent.
Borror, D. J. 1951. New Records of Maine Dragonflies (Odonata). Ent. News.
Brimley, C. S. 1903. List of dragonflies (Odonata) from North Carolina,
especially from the vicinity of Raleigh. Ent. News 14:150-157.
Brown, C. J. D. 1934. A preliminary list of Utah Odonata. Occ. Pap. Mus.
Zool., Univ. Mich. 291:1-17.


Vol. 57, No. 3, 1974

Johnson: Keys and Distribution of Calopteryx

Byers, C. F. 1931b. Dixie dragonflies collected during the summer of 1930
(Odonata). Ent. News 42:113-119.
Buchholtz, C. 1955. Eine vergleichende Ethologie der orientalischen
Calopterygiden (Odonata) als Beitrag zu ihrer systematischen Deu-
tung. Z. Tierpsychol. 12:364-386.
Calvert, P. P. 1895a. The Odonata of New York State. J. New York Ent. Soc.
Calvert, P. P. 1895b. The Odonata of Baja California, Mexico. Proc. Calif.
Acad. Sci., Series 2, 4:463-558.
Cockerell, T. D. A. 1913. The dragon-fly genus Agrion (Calopteryx) in
Colorado. Psyche 20:173-174.
Cruden, R. W. 1962. A preliminary survey of West Virginia dragonflies
(Odonata). Ent. News 73:156-160.
Donnelly, T. W. 1961. The Odonata of Washington, D. C., and vicinity. Proc.
Ent. Soc. Wash. 63:1-13.
Elrod, M. J. 1898. Iowan Odonata. Ent. News 9:7-10.
Garman, P. 1917. The Zygoptera, or damsel-flies, of Illinois. Bull. Ill. State
Lab. Nat. Hist. 12:411-587.
Garman, P. 1927. The Odonata or dragonflies of Connecticut, 331 p. Part 5 in
Guide to the insects of Connecticut. Conn. Geol. and Nat. Hist. Survey
Bull. 39.
Gibbs, R. H., and S. P. Gibbs. 1954. The Odonata of Cape Cod, Massachusetts.
J. New York Ent. Soc. 62:167-184.
Gloyd, L. K. 1951. Records of some Virginia Odonata. Ent. News 62:109-114.
Goodwin, J. T. 1968. Additions to the list of Odonata from Tennessee. J. Tenn.
Acad. Sci. 43:27.
Hagen, H. 1861. Synopsis of the Neuroptera of North America. Washington,
Smithsonian Miscel. Collections. xvii + 347 p.
Hagen, H. 1863. Abbot's Handzeichnungen in Britischen Museum und die
Neuroptera Georgiens. Ent. Zeit. (Stettin). 24:369-378.
Hagen, H. 1874. The odonate fauna of Georgia, from original drawings now in
possession of Dr. J. Le Conte, and in the British Museum. Proc. Bost.
Soc. Nat. Hist. 16:425-441.
Hagen, H. A. 1890. Synopsis of the Odonata of North America. No. 1. Psyche
Harwood, P. D. 1959. Agrion amatum (Hagen), a damselfly (Odonata) new to
Maine. Maine Field Nat. 15:54.
Harwood, P. D. 1960. Additional Notes on the Odonata (Dragonflies) of Ohio.
Ohio J. Sci. 60:341-344.
Harwood, P. D. 1972. Calopteryx angustipenne Selys in West Virginia. Proc.
W. Va. Acad. Sci. 44:89.
Howe, R. H. 1917-1921. Manual of the Odonata of New England, II. Mem.
Thoreau Mus. Nat. Hist. 1-138.
Howe, R. H. 1917. Distributional Notes on New England Odonata. Part I.
Psyche 24:45-53.
Howe, R. H. 1919. Distributional Notes on New England Odonata. Part II.
Psyche 25:106-110.
Howe, R. H. 1921. The distribution of New England Odonata. Proc. Boston
Soc. Nat. Hist. 36:105-133.
Huggins, J. R. 1927. Variations in size of Calopteryx maculata and a proposed
new subspecies. Trans. Amer. Ent. Soc. 52:355-364.
Johnson, C. 1962. Breeding behavior and oviposition in Calopteryx macula-
tur (Beauvois) (Odonata:Calopterygidae). Amer. Midl. Nat.
Johnson, C. 1972. The damselflies (Zygoptera) of Texas. Bull. Florida State
Mus., Biol. Sci. 16:55-128.


The Florida Entomologist

Johnson, C. 1973a. Distributional patterns and their interpretation in He,
taerina (Odonata:Calopterygidae). Fla. Ent. 56:24-42.
Johnson, C. 1973b. Variability, distribution, and taxonomy of Calopteryx
dimidiata (Zygoptera:Calopterygidae). Fla. Ent. 56:207-222.
Kennedy, C. H. 1915. Notes on the life history and ecology of the dragonflies
(Odonata) of Washington and Oregon. Proc. U. S. Nat. Mus.
Kennedy, C. H. 1917a. Notes on the life history and ecology of the dragonflies
(Odonata) of central California and Nevada. Proc. U. S. Nat. Mus.
Kennedy, C. H. 1917b. The Odonata of Kansas with reference to their dis-
tribution. Bull. Kansas Univ. 18(1):129-145.
Kennedy, C. H. 1918. The varieties of the dragonfly, Agrion aequabile (Say).
Can. Ent. 50:406-410.
Kimmins, D. E. 1969. A list of the type-specimens of Odonata in the British
Museum (Natural History) Part II. Bull. Br. Mus. Nat. Hist. (Ent.).
Kormondy, E. J. 1958. Catalogue of the Odonata of Michigan. Miscel. Publ.
Mus. Zool., Univ. Mich. 104:1-43.
La Rivers, I. 1940. A preliminary synopsis of the dragonflies of Nevada. Pan
Pacific Ent. 16:111-123.
Martin, R. D. C. 1939. Life histories of Agrion aequabile and Agrion macula-
turn. Ann. Ent. Soc. Amer. 32:601-618.
Montgomery, B. E. 1935. Records of Indiana dragonflies, VIII, 1934. Proc. Ind.
Acad. Sci. 44:231-235.
Montgomery, B. E. 1937. Records of Indiana dragonflies, IX, 1935-1936. Proc.
Ind. Acad. Sci. 46:203-210.
Montgomery, B. E. 1940. The Odonata of South Carolina. Elisha Mitchell Sci.
Soc. 56:283-301.
Montgomery, B. E. 1941. Records of Indiana dragonflies, X, 1937-1940. Proc.
Ind. Acad. Sci. 50:229-241.
Montgomery, B. E. 1943. Records of Ohio dragonflies. Ohio J. Sci. 43:267-270.
Montgomery, B. E. 1947. The distribution and relative seasonal abundance of
Indiana species of five families of dragonflies (Odonata,
Calopterygidae, Petaluridae, Cordulegasteridae, Gomphidae and
Aeshnidae). Proc. Ind. Acad. Sci. 56:163-169.
Montgomery, B. E. 1951. Notes and records of Indiana Odonata, 1941-1950.
Proc. Ind. Acad. Sci. 60:205-210.
Montgomery, B. E. 1953. Notes and records of Indiana Odonata, 1951-1952.
Proc. Ind. Acad. Sci. 62:200-202.
Montgomery, B. E. 1955. Notes and records of Indiana Odonata, 1953-1954.
Proc. Ind. Acad. Sci. 64:131-135.
Montgomery, B. E. 1971. Records and observations of Indiana Odonata. Proc.
Ind. Acad. Sci. 80:253-263.
Murphey, E. E. 1945. Historical narrative. In: Birds of Georgia, a preliminary
check-list and bibliography of Georgia ornithology. Compiled by E. R.
'Greene etc. Univ. Georgia Press, Athens. 111 p.
Muttkowski, R. A. 1908. Review of the dragonflies of Wisconsin. Bull. Wisc.
Nat. Hist. Soc. 6:57-126.
Muttkowski, R. A. 1909. A summer's insect collecting. Bull. Wisc. Nat. Hist.
Soc. 6:164-169.
Muttkowski, R. A. 1910. Catalog of the Odonata of North America. Bull. Pub.
Mus., Milwaukee. 1:1-207.
Needham, J. G. 1903. Aquatic insects in New York State. Part 3. Life histories
of Odonata, Suborder Zygoptera, damsel flies. Bull. N. Y. State Mus.
Needham, J. G. 1928. A list of insects of New York. Order Odonata. Cornell
Univ. Agr. Exp. Station Mem. 101:45-56.

Vol. 57, No. 3, 1974

Johnson: Keys and Distribution of Calopteryx

Needham, J. G., and H. Heywood. 1929. A handbook of the dragonflies of
North America. Springfield, Ill. VIII + 378 p.
Newell, R. L. 1970. Checklist of some aquatic insects from Montana. Proc.
Montana Acad. Sci. 30:45-56.
Pajunen, V. I. 1966. Aggressive behaviour and territoriality in a population of
Calopteryx virgo L. (Odon., Calopterygidae). Ann. Zoo. Fennici
Resner, P. L. 1970. An annotated check list of the dragonflies and damselflies
(Odonata) of Kentucky. Trans. Kentucky Acad. Sci. 3:32-44.
Roback, S. S., and Minter J. Westfall, Jr. 1967. New records of Odonata
nymphs from the United States and Canada with water quality data.
Trans. Amer. Ent. Soc. 93:101-124.
Robert, Adrien. 1963. Les Libellules du Quebec. Service de la Fauna Bull. 1.
Minister de la Chasse et de la Peche, Quebec. viii + 223 p.
Root, F. M. 1923. Notes on Zygoptera (Odonata) from Maryland, with a
description of Enallagma pallidum n. sp. Ent. News 34:200-204.
Root, F. M. 1924. Notes on dragonflies (Odonata) from Lee County, Georgia,
with a description of Enallagma dubium new species. Ent. News
Seeman, T. M. 1927. Dragonflies, mayflies, and stoneflies of southern Califor-
nia. J. Ent. and Zool. Pomona Coll. 19:1-69.
Selys Longchamps, Le Baron Edmond De. 1853. Synopsis des Calopterygines.
Brussells, Hayez. 73 p.
Selys Longchamps, Le Baron Edmond De. 1854. Monographie des
Calopterygines. Mem. Soc. roy. des Sciences de Liege. Brussells and
Leipzig. xii + 291 p.
Smith, R. F., and A. E. Pritchard. 1956. Chapter 4, Odonata. In: Aquatic
insects of California, ed. by R. L. Usinger. Univ. Calif. Press x + 508 pp.
Trogdon, R. P. 1961. A survey of the adult Odonata of Tennessee. Unpublished
Ph.D. diss. submitted to the Univ. of Tennessee. 1961.
Walker, E. M. 1953. The Odonata of Canada and Alaska. Vol. 1. Univ. Toronto
Press, Toronto. xi + 292 p.
Wells, Lloyd. 1917. Odonata of Iowa. Proc. Iowa Acad. Sci. 24:327-333.
Whedon, A. D. 1914. Preliminary notes on the Odonata of Southern
Minnesota. Minn. State Ent. Report for 1914:77-103.
White III, H. B., and W. J. Morse. 1973. Odonata (Dragonflies) of New
Hampshire: an annotated list. Res. Rep. 30, N. H. Agric. Exp. St.,
Durham. v + 46 p.
Williamson, E. B. 1899. Calopteryx angustipennis Selys in western Pennsyl-
vania. Ent. News 10:199-200.
Williamson, E. B. 1900. The dragonflies of Indiana. Rep. Indiana State
Geologist. 233-333 pp. index and glossary, 1003-1010 pp.
Williamson, E. B. 1903. The dragonflies (Odonata) of Tennessee, with a few
records for Virginia and Alabama. Ent. News 14:221-229.
Williamson, E. B. 1904. The dragonflies (Odonata) of Burma and Lower
Siam-I. Subfamily Calopteryginae. Proc. U. S. Nat. Mus. 28:165-187.
Williamson, E. B. 1917. An annotated list of the Odonata of Indiana. Univ.
Mich. Mus. Zool., Misc. Publ. 2:1-12.
Williamson, E. B. 1923. Odonatological results of an auto trip across Indiana,
Kentucky, and Tennessee. Ent. News 24:6-9.
Williamson, E. B. 1932. Dragonflies collected in Missouri. Occ. Pap. Mus.
Zool., Univ. Mich. 240:1-40.
Wilson, C. B. 1909. Dragonflies of the Mississippi Valley collected during the
Pearl Mussle Investigations on the Mississippi River, July and August,
1907. Proc. U. S. Nat. Mus. 36:653-671.
Wilson, C. B. 1912. Dragonflies of the Cumberland Valley in Kentucky and
Tennessee. Proc. U. S. Nat. Mus. 43:189-200.

The Florida Entomologist

Woodruff, L. B. 1914. Some dragonflies of a Connecticut brook. J. New York.
Ent. Soc. 22:154-159.
Wright, M. 1943. Dragonflies collected in the vicinity of Florala, Alabama.
Fla. Ent. 26:30-31; 49-51.

The following entry, received from R. W. Garrison subsequent to writing the
above paper, increases the scanty California distribution for Calopteryx
Mendocino Co., 15-16 June 1972. CIS Collection, Univ. of Cal., Berkeley.


Specializing in 4Boohs and cpubl'cations

Storter Printing Co.



Vol. 57, No. 3, 1974

The Florida Entomologist

Department of Entomology and Economic Zoology,
Clemson University, Clemson, South Carolina 29631


Stiretrus anchorago (F.) was collected from S. C. soybean fields and reared
at 3 constant temperatures. Larvae of Epilachna varivestis Mulsant were
used as prey for S. anchorago in life history studies.
Female adults lived an average of 46.0, 29.6, and 12.6 days; males 38.2, 24.8,
and 22.4 days at constant temperatures of 18.3, 26.7, and 32.20 C, respectively.
Mean numbers of eggs per female were 12.6, 57.3, and 15.0 at 18.3, 26.7, and
32.20 C, respectively.
The immature stages (eggs and instars) completed development in 42.8,
24.6, and 22.6 days at 18.3, 26.7, and 32.20 C, respectively.
Eggs laid by field collected females and held at 26.70 C were more viable
(88.5%) than eggs from laboratory reared females (19.0%). Only infertile eggs
were produced by females fed Galleria mellonella (L.) larvae exclusively.

Stiretrus anchorago (F.) has been reported to be the most numerous
pentatomid attacking the Mexican bean beetle, Epilachna varivestis Mul-
sant, in Mexico and the southern part of the United States (Howard 1936). It
has also been found in the central, Middle Atlantic, and New England states.
Plummer and Landis (1932) found S. anchorago widely distributed in Mexico.
Blatchley (1926) listed S. anchorago and S. fimbriatus (Say) as the only 2
species of Stiretrus found in the eastern U. S. The 2 species are easily separated
as S. anchorago is black and red, black and orange-yellow, or violet while S.
fimbriatus is brown-bronzed and dull yellow (Blatchley 1926).
Getting and Yonke (1971) described the immature stages and biology of S.
fimbriatus, and Howard (1936) briefly mentioned development and feeding of
S. anchorago.
In order to better understand the biology and predatory potential of S.
anchorago, laboratory studies were conducted from July until December,
S. anchorago nymphs and adults were collected from soybean fields at
Clemson University, Edisto Experiment Station, Blackville, S. C., and were
returned to the laboratory for life history studies. Field collected adults were
maintained in pint cardboard containers with screened lids and were fed E.
varivestis larvae. Water was provided in a moist cotton ball placed on top of

I Technical contribution number 1140. Published by permission of the Director, South Carolina
Agricultural Experiment Station.

Vol. 57, No. 3, 1974

The Florida Entomologist

the screened lid. Approximately 5 pairs of adults were held per container. The
stock colony was kept in a rearing room at 26.7 +1.10 C, 60+10% RH, and a
photoperiod of LD 14:10. Additional environmental chambers were set at
18.3- +1.1 and 32.2 + 1.10 C, 60 +.10% RH and a photoperiod of LD 14:10.
Longevity and fecundity of adult S. anchorago were studied by placing a
single pair of newly emerged adults in a plastic dish (7.8 cm diam X 4.6 cm).
The dish contained a 2-ml vial glued in a vertical position to the dish bottom
and filled with water which supported a single lima bean leaf. The leaf
provided moisture for the pentatomids and foliage for the four 4th-stage
Mexican bean beetle larvae which were provided daily as food. The bottom of
each dish was covered with a thin layer of cotton to provide better footing for
S. anchorago. Data were recorded every 24 hr.
Duration and survival of the nymphal stages were determined by placing
individual nymphs in plastic containers as previously described. Nymphs were
fed either 3rd or 4th-stage E. varivestis.
Duration of the egg stage was determined by placing individual egg masses
in 100 x 15 mm plastic petri dishes which contained a moist cotton ball. The
petri dish was then placed in the appropriate temperature regime and in-
spected daily until eclosion.

Females lived slightly longer than males at 18.3 and 26.70 C (Table 1).
However, males appeared to be more tolerant of 32.2 C. Longevity of both
sexes was extended with decreasing temperatures. Extreme temperatures
(18.3 and 32.20 C) reduced egg production per female by ca. 75% below
production at 26.70 C, although the number of days between oviposition and
the preoviposition period was reduced as temperature was increased. S.
anchorago produced only about 1/6 as many eggs as reported for Podisus
maculiventris (Say) (Warren and Wallis 1971), another commonly-occurring
predaceous pentatomid in S. C. soybeans.
Adults fed voraciously on E. varivestis larvae, but were reluctant to feed
on larvae of Galleria mellonella L. When only G. mellonella larvae were
supplied as food, S. anchorago produced very few eggs, none of which were
viable. We also found that if starved, adults and immatures fed on larvae of
the soybean looper, Pseudoplusia includes (Walker). This apparent
preference for Mexican bean beetle larvae may partially explain the low
relative abundance of S. anchorago although their numbers were greater late
in the season which coincided with an increase in the numbers of Mexican
bean beetles.
Male and female adults differed markedly in their weights, with males
averaging 47 mg and females 74 mg (20 replicates each). Males were also easily
recognized by depressed pubescent patches on either side of sternal segments
Eggs produced by laboratory-reared females were less viable than eggs
produced by field-collected females (Table 2). The highest temperature (32.20
C) reduced egg hatch as only 1.4% hatched while 19.0 and 25.0% hatched when
held at 26.7 and 18.30 C, respectively.
First instars did not require food to develop to the second instar, but soon
died if water was not provided. This was also noted by Oetting and Yonke
(1971) in their study of S. fimbriatus and by Mukerji and LeRoux (1965) with
P. maculiventris.


Vol. 57, No. 3, 1974

Waddill and Shepard: Biology of Stiretrus anchorage 251

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The Florida Entomologist


Source Temp. N % Mean time to eclosion
of (oC) Hatch SD (days)

18.3 80 25.0 8.2+0.7
Lab 26.7 100 19.0 6.3 0.5
Reared 32.2 68 1.4 4.0 + 0.0
Collected 26.7 391 88.5 6.2+0.6

Duration and percent survival of the 5 instars are presented in Table 3.
First instars had a higher survival rate (90%) than 5th instars (77.7%) at 18.30
C. However, at 32.2 C, 5th instar survivorship was considerably higher
(63.6%) than that for 1st instars (30.4%).
Total duration of the nymphal stages was 34.6, 18.4, and 18.6 days for the
temperatures 18.3, 26.7, and 32.20 C, respectively (Table 3). The time spent in
each immature stage agrees in general with the report of Howard (1936),
although his experiments were not conducted at constant temperatures. Time
required for development of S. anchorago was also similar to that of S.
fimbriatus (Oetting and Yonke 1971).
Further studies on prey consumption and preference will aid in under-
standing the role of S. anchorago as a predator in the soybean agroecosys-


Blatchley, W. S. 1926. Heteroptera or true bugs of Eastern North America.
Nature, Indianapolis. 116 p.
Howard, N. F. 1936. Parasites and predators of the Mexican bean beetle in the
United States. USDA Circ. 418:1-12.
Mukerji, M. K., and E. J. LeRoux. 1965. Laboratory rearing of a Quebec strain
of the pentatomid predator, Podisus maculiventris (Say) (Hemiptera:
Pentatomidae). Phytoprotection 46(1):40-60.
Getting, R. D., and T. R. Yonke. 1971. Immature stages and biology of Podisus
placidus and Stiretrus fimbriatus (Hemiptera: Pentatomidae). Can.
Ent. 103:1505-16.
Plummer, C. C., and B. J. Landis. 1932. Records of some insects predaceous on
Epilachna corrupt Muls. in Mexico. Ann. Ent. Soc. Amer. 25:695-708.
Warren, L. 0., and G. Wallis. 1971. Biology of the spined soldier bug, Po-
disus maculiventris (Hemiptera: Pentatomidae). J. Georgia Ent.
Soc. 6(2):109-16.


Vol. 57, No. 3, 1974

Waddill and Shepard: Biology of Stiretrus anchorago 253


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The Florida Entomologist



Department of Biological Sciences,
Illinois State University, Normal, Illinois 61761

The Echmepteryx hageni complex in Florida includes 3 species. Two are
described as new and a full description of E. hageni (Packard) is presented. In
Florida, E. young n. sp. seems to be the only bisexual member of the complex.
Males of E. intermedia n. sp. are unknown, and those of E. hageni are
extremely rare. In southern Florida E. young occurs in most plant com-
munities dominated by trees or shrubs but where its range overlaps that of E.
hageni, it is restricted almost entirely to the sand scrub communities and
occurs primarily on 2 species of pine.

The type species, Echmepteryx hageni (Packard), of a largely tropical
genus, occurs commonly on tree trunks and branches throughout the eastern
United States and southeastern Canada. Forms, presumably all distinct
species, very closely allied to it occur in Florida, most islands of the Caribbean,
southeastern Mexico, and Belize. These presumed species plus E. hageni
constitute the Echmepteryx hageni complex.
In Florida and the southeastern corner of Georgia this complex consists of
3 discrete species. Two are allopatric with respect to each other, while the
third broadly overlaps the ranges of the other 2.
The purpose of this paper is to describe 2 of these species as new, to present
an augmented description of E. hageni, and to discuss ecological and
phylogenetic relationships among these species.
In all members of the complex described here, the face is marked with a
singular pattern (Fig. 1, 4, 7) always involving an arched mark in each parietal
region over the lateral ocellus. The lacinial tip is tridentate (Fig. 20) and the
pretarsal claw (Fig. 21) bears a large preapical denticle, several minute den-
ticles at the ends of striae in the middle region, and a basal acuminate bristle.
The female subgenital plate has a small triangular pigmented area with apex
of the triangle anterior; the area is setose with 4 setae longer than the others
on its posterior margin. The sense cushion of the paraproct is diffuse and
includes 6 trichobothria and 1 long seta with no basal floret. Female
gonapophyses, as in all members of the genus, are reduced to the setose third
valvulae, which are elongate, slender, and joined to each other by membrane
along the mid-line. The spermathecal duct from its external orifice for a short
distance back is surrounded by a sclerotized collar. Proportions of this collar
differ in the different species.

Contribution No. 306. Bureau of Entomology, Division of Plant Industry, Florida Department
of Agriculture and Consumer Services, Gainesville, Fla. 32602.
Research Associate, Florida State Collection of Arthropods, Fla. Dep. Agr. and Cons. Serv.,

Vol. 57, No. 3, 1974







I, L/)




Fig. 1-9 Echmepteryx hageni complex species: 1) E. hageni (Packard)
female, facial markings (scales in mm); 2) E. hageni (Packard) female, fore-
wing; 3) E. hageni (Packard) female, anterior tibia; 4) E. young n. sp. fe-
male, facial marking, scale of Fig. 1; 5) E. young n. sp. female, forewing,
scale of Fig. 2; 6) E. young n. sp. female, anterior tibia, scale of Fig. 3;
7) E. intermedia n. sp. female, facial markings, scale of Fig. 1; 8) E. inter-
media n. sp. female, forewing, scale of Fig. 2; 9) E. intermedia n. sp. fe-
male, anterior tibia, scale of Fig. 3.



O i



12 Q05

Fig. 10-18 Echmepteryx hageni complex species: 10) E. hageni (Packard)
female, left valvula; 11) E. hageni (Packard) female, collar of spermathecal
duct; 12) E. hageni (Packard) female, distal segment of maxillary palpus; 13)
E. young n. sp. female, left valvula, scale of Fig. 10; 14) E. young n. sp. female,
collar of spermathecal duct, scale of Fig. 11; 15) E. young n. sp. female, distal
segment of maxillary palpus, scale of Fig. 12; 16) E. intermedia n. sp. female,
left valvula, scale of Fig. 10; 17) E. intermedia n. sp. female, collar of sper-
mathecal duct, scale of Fig. 11; 18) E. intermedia n. sp. female, distal segment
of maxillary palpus, scale of Fig. 12.

The Florida Entomologist

Material examined consisted of 250 adult specimens and 140 nymphs, the
latter determined to species by their facial pattern. Of these, 91 adults and 23
nymphs represent E. hageni, 123 adults and 72 nymphs represent E. young,
and 36 adults and 45 nymphs represent E. intermidia. With exception of 1
locality in southeastern Georgia near the Florida line, all material is from
Measurements are stated in microns and have an error of +0.7754.
Abbreviations for structures measured are as follows: Fw = forewing length; f,,
f= lengths of first and second flagellomeres; F=posterior femur length;
T= posterior tibial length; t,, t,, t,= lengths of first, second, and third tar-
someres. The micrometer unit used in constructing the scatter diagram (Fig.
23) of relationship of forewing length to greatest forewing width is
approximately 18.3p and is not converted to metric units in the diagram.

Echmepteryx hageni (Packard)
Amphientomum hageni Packard, 1870:405.
Echmepteryx agilis Aaron, 1886:17.
Echmepteryx hageni (Packard), Enderlein 1906:104.
Diagnosis.-Parietal arched marks of face including a narrow band on each
side close to and paralleling margin of compound eye (Fig. 1). Forewing rela-
tively broad (Fig. 2, 23). Third valvula long and slender (Fig. 10).
Color (in alcohol; sexes similar).-Compound eye of fresh specimen pale
grayish-green with a dark brown band running from region of antero-ventral
eye margin nearest antennal socket postero-dorsally, not reaching posterior
margin. Compound eye black on specimens in alcohol several months or more.
Face marked (Fig. 1), with dark areas dark brown, pale areas dull creamy
white to pale brown. Thorax dorsally clothed in dark brown scales, the cuticle
under these medium brown. Thoracic pro- and mesopleura dark brown;
remainder of thorax pale to medium brown. Legs from coxa to middle of each
femur pale to medium brown. Distal half of each femur darker brown. Each
tibia banded in pattern of Fig. 3 with dull creamy white and dark brown. Each
tarsus with dark brown basal band, fading distally, occupying basal third of t,;
remainder dull creamy white. Forewings clothed in dark brown scales, the
membrane medium brown except colorless in distal fifth. Preclunial ab-
dominal segments sparsely clothed in medium brown scales; the cuticle (or
subcuticular pigment?) pale brown. Clunium, epiproct, paraprocts, and ex-
ternal genitalia dark brown.
General morphological features.-Head in frontal view with posterior
margin approximately straight, slightly depressed at mid-line. Fourth seg-
ments of maxillary palpus (Fig. 12) relatively slightly dilated toward apex.
Male external genitalia.-Hypandrium smoothly rounded posteriorly, se-
tose over its entire margin except on lateral margins. Phallosome (Fig. 19) of
general type seen in E. madagascariensis (Kolbe) and E. terricolis Badonnel.
External parameres long and slender, distally enclosing at least in part the
internal parameres; externals bearing a few pores near their tips, tapering at
their tips. Internal parameres elongate, slender, hollowed internally, also
bearing pores. At bases of internal parameres a pair of bulb-like structures,
and basal to these a cup-shaped region divided medially.
Female genitalia.-Third valvula as described in diagnosis. Collar of sper-
mathecal duct (Fig. 11) elongate and slender.


Vol. 57, No. 3, 1974

Mockford: The Echmepteryx hageni Complex

Distribution.-Eastern United States and southeastern Canada west to
central Iowa, southeastern Kansas, and eastern Texas.
Florida records (For all records in this paper unless otherwise stated,
I was the collector).-Alachua Co.: Cross Creek Hammock, 15 Nov. 1952,
beating red maples (Acer rubrum) along creek, 1 female; Newnan's Lake,
2 May and 11 June 1952, tree trunks, vines, and branches, 2 females.
Citrus Co.: 4 mi. n. Crystal River on Highway 19, 15 April 1965, trees and
shrubs in hammock, 1 female, 2 ny. Clay Co.: Goldhead Branch State
Park, 20-22 Oct. 1973, beating turkey oak (Quercus laevis) and live oak
(Quercus virginiana) in sand scrub, 16 females, 3 ny.; 3 mi. ne. Keystone
Heights on State Highway 21, 22 Oct. 1973, beating long-leaf pine (Pinus
australis), 2 females, 3 ny.; Magnolia Lake State Recreation Area, 22 Oct.
1973, beating long-leaf pine, 6 females, 6 ny. Columbia Co.: O'leno State
Park, 30 Oct.-14 Nov. 1953, trunks and branches of river birch (Betula
nigra), dogwood (Cornus sp.), and bald cypress (Taxodium distichum)
along Santa Fe River, 5 females, 1 ny.; same loc., 23 Oct. 1973, beating
oaks in sand scrub, 7 females, 3 ny.; beating long-leaf pine in sand scrub,
2 females. Franklin Co.: 1 mi. s. Ochlockonee River on Highway 98, 25
Oct. 1973, beating long-leaf pine, 1 female. Leon Co.: Apalachicola Na-
tional Forest, 10 mi. w. Tallahassee, July 1971, beating live oak, slash
pine (Pinus elliottii), and holly, 1 female, coll. L. and J. Brown; 6 mi. s.
Tallahassee on State Highway 369, 25 Oct. 1973, beating turkey oak and
live oak in sand scrub, 9 females. Liberty Co.: Torreya State Park, 22
August 1951, trunk of beech (Fagus grandifolia), 1 female; same loc., 23
June 1956, 1 ny., coll. P. Kannowski. Nassau Co.: 12 mi. ne. Bryceville on
Highway 301, 17 Oct. 1973, beating slash pine and loblolly pine (Pinus
taeda), 7 females, 1 ny. Okaloosa Co.: Blackwater River State Forest (Na-
ture Center), 27 Oct. 1973, beating white cedar (Chamaecyparis thyoides),
6 females. Santa Rosa Co.: Blackwater River State Forest (Fish
Hatchery), 28 Oct. 1973, beating pine and oaks, 8 females. Union Co.:
Itchetucknee Springs, 20 June 1953, beating junipers (Juniperus sp.), lob-
lolly pine, and broad-leaved trees and shrubs, 3 females, 2 ny. Wakulla
Co.: 6 mi. n. Crawfordsville on State Highway 369, 25 Oct. 1973, beating
broad-leaved trees and shrubs, 2 females, 2 mi. n. Ochlockonee River on
Highway 98, beating long-leaf pine, 3 females, 1 ny.; 9 mi. n. Sopchoppy
on State Highway 375, 23 August 1951, sweeping dead branches in bay-
head, 1 male, 1 female. Walton Co.: Villa Tasso, 27 Oct. 1973, beating
sand pine (Pinus clausa), turkey oak, and live oak in sand scrub, 4 females.
Georgia.-Camden Co.: Crooked River State Park, 16 Oct. 1973, beat-
ing scrub li e oak (Quercus geminata), and long-leaf pine, 2 females.

Echmepteryx young, n.sp.
Diagnosis.-Parietal arched marks of face lacking a narrow band on each
side paralleling margin of compound eye (Fig. 4). Forewing relatively slender
(Figs. 5, 23). Third valvula (Fig. 13) relatively short and stubby.
Color (in alcohol; sexes similar).-Compound eye in fresh specimen pale
grayish-green with a dark brown band of irregular width from just above
antero-ventral margin toward postero-dorsal margin, ending before reaching
latter margin, and a large dark brown blotch in region of eye bordering gena.
Compound eyes black in specimens in alcohol several months or more. Face
marked as in Fig. 4, with dark areas dark brown, pale areas dull creamy white
to pale brown. Thorax dorsally clothed in medium brown, shining scales, the
cuticle under these medium brown. A broad band through pro- and
mesopleura dark brown; remainder of thorax creamy white. Coxae mottled

The Florida Entomologist

dusky brown to creamy white; femora creamy white except medium brown
distally. Tibiae banded as in Fig. 6 with dull creamy white and dark brown, the
dark brown bands relatively narrower than in E. hageni. Each tarsus with
dark brown basal band, fading distally, occupying basal third of t,. Forewings
(Fig. 5) spottily clothed in dark brown scales, the membrane variegated brown
to clear. Preclunial abdominal segments sparsely clothed in medium brown
scales; subcuticular pigment pale brown, darker on sides. Clunium, epiproct,
paraprocts, and external genitalia dark to pale brown.
Measurements.-Table I.
General morphological features.-Head in frontal view (Fig. 4) with
posterior margin slightly indented at mid-line, from there arching slightly up
on each side and down to compound eye margin. Fourth segment of maxillary
palpus (Fig. 15) decidedly more dilated toward its apex than in E. hageni.
Male external genitalia.-Hypandrium as described for E. hageni.
Phallosome (Fig. 22) in general as described for E. hageni but differing as
follows: external parameres each bearing a decided transverse crease near its
tip; tip more bent inward; internal parameres distad of the 2 bulbous struc-
tures decidedly shorter; each internal paramere decidedly constricted in a

S19 22




Fig. 19-22 Echmepteryx hageni (Packard) and E. young n. sp.: 19) E.
hageni (Packard) male, phallosome; 20) E. hageni (Packard) male, lacinial
tip; 21) E. hageni (Packard) female, pretarsal claw; 22) E. young n. sp. male,
phallosome, twice scale of Fig. 19.

Vol. 57, No. 3, 1974

Mockford: The Echmepteryx hageni Complex

short region just distad of the bulbous structures; the bulbous structures
narrower; the divided cup-shaped region basal to the two bulbous structures
more diffuse.
Female genitalia.-Third valvula as described in diagnosis. Collar of sper-
mathecal duct (Fig. 14) relatively short.
Type locality.-Florida: Sarasota Co.: Myakka River State Park, 30
August 1951, on trunks of cabbage palms (Sabal palmetto); holotype male,
allotype female, 1 male, and 6 female paratypes. The type material is in
the author's collection.
Records.-Florida: Alachua Co.: Newnan's Lake, 11 Oct. 1953, on side
of wooden shed, 1 female. Clay Co.: Goldhead Branch State Park, 20-21
Oct. 1973, beating long-leaf pine and turkey oak in sand scrub, 1 male,
4 females; Magnolia Lake State Recreation area, 22 Oct. 1973, beating
long-leaf pine, 3 females. Collier Co.: State Highway 846 2 mi. w. Naples
Park Beach, 29 Nov. 1970, beating vegetation, 2 males, 7 females, 6 ny.
Franklin Co.: 1 mi. s. Ochlockonee River on Highway 98, 25 Oct. 1973,
beating long-leaf pine, 2 males, 1 female, 1 ny.; 4 mi. sw. Ochlockonee
River on Highway 98, 25 Oct. 1973, beating sand pine, 1 male, 8 females.
Indian River Co.: Vero Beach, 18 April 1954, beating cabbage palm, 1
female, 5 ny. Manatee Co.: Highway 301, 3 mi. s. Parrish, 4 Dec. 1970,
beating pines and cabbage palms, 1 male, 1 female, 3 ny. Monroe Co.: Dry
Tortugas: Loggerhead Key, 11 Jan. 1962, on geiger tree (Cordia sebes-
tena), 2 females, coll. H. A. Denmark; same loc., 5 June 1962, in black
light trap, 1 male, 1 female, coll. H. V. Weems Jr. and R. W. Woodruff;
Inner Keys: Island E-9, Calusa Keys near Upper Matecumbe Key, 27
Feb. 1967, 2 females, 1 ny., coll. E. O. Wilson; Long Key, 13 April 1953, on
thatch palm (Thrinax sp.) leaves, 1 male; Outer Keys: Island Cr-1, Crane
Keys, 18 July 1969, 3 females, 9 ny., coll. D. Simberloff; Island J-1 near
Johnston Key, 13 August 1969, 6 males, 5 females, 14 ny., coll. D. Simber-
loff; Island Mud-1, Mud Keys, 4 July 1969, 8 males, 13 females, 15 ny.,
coll. D. Simberloff; Island Mud-2, Mud Keys, 24 July 1969, 1 male, 7 fe-
males, 16 ny., coll. D. Simberloff; Island E-2, Snipe Keys, 8 Dec. 1967, 1
female, coll. D. Simberloff. Palm Beach Co.: Highway 441 w. Boynton
Beach, 3 Dec. 1970, beating slash pines, 1 female. Wakulla Co.: 2 mi. n.
Ochlockonee River on Highway 98, 25 Oct. 1973, beating long-leaf pine,
1 male.
Georgia.-Camden Co.: Crooked River State Park, 16-17 Oct. 1973,
beating long-leaf pine and scrub live oak, 15 males, 12 females, 2 ny.
I take pleasure in naming this species for Dr. Frank N. Young, who in-
troduced me to the natural history of Florida.

Echmepteryx intermedia, n. sp.
Diagnosis.-Parietal arched marks of face (Fig. 7) lacking a narrow band on
each side paralleling margin of compound eye. Forewing relatively slender, of
intermediate width between those of E. hageni and E. young (Fig. 8, 23).
Third valvula (Fig. 16) shorter than in E. hageni, somewhat longer and more
slender than in E. young.
Color (in alcohol).-Compound eyes black in specimens in alcohol several
months or more. Face marked as in Fig. 7, with dark areas dark brown, pale
areas dull creamy white. Generally each parietal arched mark with a pale spot
in its broadest part. A narrow brown band on posterior surface of head behind
parietal mark. Thorax dorsally at least partially clothed in medium brown
Measurements.-Table I.

The Florida Entomologist

Vol. 57, No. 3, 1974

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Mockford: The Echmepteryx hageni Complex

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The Florida Entomologist


0 08 o o
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600 700 800

Fig. 23. Forewing length (horizontal axis) plotted against greatest fore-
wing width (vertical axis) for females of 3 species of Echmepteryx: tri-
angles = E. young n. sp.; diamonds = E. intermedia n. sp.; circles = E. hageni

scales, the cuticle under these medium brown. Thoracic pleura and legs as
described for E. young, except dark tibial bands slightly wider (Fig. 9). Tarsi as
described for E. young. Forewings clothed in brown scales (largely rubbed off
in specimens on hand); long marginal scales in alternating patches of dark and
pale around distal margin of wing. Forewing membrane mostly pale brown,
with clear basal and distal regions as indicated in Fig. 8. Preclunial abdominal
segments sparsely clothed in medium brown scales; subcuticular pigment
creamy white, somewhat darker on sides. Clunium, epiproct, paraprocts, and
external genitalia dark to medium brown.
General morphological featu-res.-Head in frontal view (Fig. 7) with
posterior margin slightly indented at mid-line, approximately straight on
either side. Fourth segment of maxillary palpus (Fig. 18) more dilated distally
than in E. hageni, its distal margin more oblique than in E. young.
Female genitalia.-Third valvula as described in diagnosis. Collar of sper-
mathecal duct (Fig. 17) relatively short and slender.
Type locality.-Florida: Palm Beach Co.: Highway 441 w. Boynton
Beach, 3 Dec. 1970, beating slash pines, holotype female, 6 female para-
types, and 5 ny. The types are in my collection.
RECORDS.-Florida: Collier Co.: Collier Seminole State Park, 23 Feb.
1956, on royal palm (Roystonea regia) trunk in hammock, 1 female; 2 mi.


Vol. 57, No. 3, 1974

Mockford: The Echmepteryx hageni Complex

w. Naples Park Beach, 29 Nov. 1970, beating vegetation, 6 females, 6 ny.;
Tamiami Trail 2 mi. e. Ochopee, 23 Feb. 1956, 1 female, 1 ny. Hendry Co.:
LaBelle, 4 Feb. 1953, beating oaks, 3 females, 5 ny.; State Highway 833,
16 April 1954, beating trees and shrubs in subtropical hammock, 2 fe-
males, 9 ny. Manatee Co.: Highway 301, 3 mi. s. Parrish, 4 Dec. 1970,
beating slash pine and live oak, 6 females, 14 ny. Monroe Co.: Key Largo,
21 Feb. 1956, trunk of gumbo limbo (Bursera simaruba), 1 female. Polk
Co.: 2 mi. n. Lake Wales on Highway 27, 25 Nov. 1961, beating live oaks,
1 female, coll. E. L. Mockford and R. O. Rilett. Sarasota Co.: Myakka
River State Park, 30-31 August 1951, on trunks of cabbage palm in ham-
mock, 2 females,1 ny.; same loc., 12 April 1952, on trunks and leaves of
cabbage palm, 2 females, 3 ny.; same loc. 3 Feb. 1953, beating cabbage
palms and live oaks, 3 females; same loc., 6-7 April 1966, beating oak
branches, 1 female, 1 ny.
Discussion.-In an earlier paper (Mockford, 1971) I considered the Florida
species of this complex to be a single species. I noted the existence of the 3
forms which, without stating it, I thought were geographic races. I also noted
the regular occurrence of males of "form B" (E. young) in the southern half of
the Florida Peninsula. Data gathered since that publication strongly suggest
the new interpretations given here. E. young occurs throughout most of
Florida (Fig. 24) and is apparently bisexual throughout its range. It is sym-
patric with E. intermedia and overlaps the southern edge of the range of E.
hageni. There is no evidence for its hybridization with either of the other 2
species, both of which seem to be parthenogenetic. Considering the absence of
hybridization as well as its morphological differences, it must be regarded as a
distinct species from either of the other 2.
Parthenogenesis has already been shown for E. hageni (Mockford 1971)
and, to my knowledge, only a single male has been taken in Florida. This male
was taken on my first collecting trip to Florida in 1951. Absence of males in
any subsequent collections suggests that there is no bisexual population of the
species in the state.
Parthenogenesis in E. intermedia must be verified by additional collecting
and laboratory rearing, but absence of males to date is certainly suggestive.
E. hageni in Florida and E. intermedia appear to be allopatric
agamospecies. There is a collecting hiatus between their known ranges, and
filling in of the hiatus will be important in testing this conclusion.
Morphologically, the 2 are completely separable and at least as distinct as
some sympatric pairs of biospecies known in the Psocoptera.
E. young shows a notable narrowing of its habitat in northern Florida
where it overlaps with E. hageni. South of the range of E. hageni, it has been
taken on trunks and leaves of cabbage palm, leaves of thatch palm,
mangroves, geiger tree, and slash pine. Thus, it occurs in the major tree- and
shrub-dominated vegetation types of southern Florida: hardwood hammocks,
mangrove swamp, pine flatwoods, and coastal scrub. Within the range of E.
hageni, E. young has been found almost entirely in sand scrub, where it
occurs primarily on long-leaf pine and sand pine, and to a lesser extent on
turkey oak and live oak. In northern Florida it has never been taken in pine
flatwoods, and only a single specimen has been found in a hardwoods ham-
mock. In Florida, E. hageni occurs in all tree-dominated vegetation types
within its range, but in sand scrub it appears to be more abundant on oaks
than on long-leaf pine, and it has never been taken on sand pine. Habitat
restriction of E. young in northern Florida may have resulted from competi-
tion with E. hageni.


The Florida Entomologist

O hageni
A young
0 intermedia

Fig. 24. Distribution pattern of 3 species of Echmepteryx in Florida.
Superimposed symbols and pairs of symbols connected by an arrow indi-
cate sympatry.

The majority of records from the Florida Keys resulted from material
contributed by Dr. E. O. Wilson of Harvard University and Dr. D. Simberloff


Vol. 57, No. 3, 1974


Mockford: The Echmepteryx hageni Complex

of Florida State University. Four specimens, including the only material on
hand from the Dry Tortugas, were borrowed from the Florida State Collection
of Arthropods, Gainesville, through the curator, Dr. Howard V. Weems, Jr.
Specimens also were contributed by Dr. and Mrs. Lauren Brown and Dr. R. O.
Rilett of Illinois State University, Dr. P. Kannowski of the University of
North Dakota, and Mr. A. Manzano, Normal, Illinois. I acknowledge with
thanks the loans and gifts of material from the above named individuals and


Aaron, S. F. 1886. On some new Psocidae. Proc. Acad. Nat. Sci. Phila. 38:13-18,
pl. 1.
Enderlein, G. 1906. Einige Notizen zur Kenntniss der Copeognathen Nor-
damerikas. Stett. Ent. Zeit. 67:317-320.
Mockford, E. L. 1971. Parthenogenesis in Psocids (Insecta: Psocoptera). Am.
Zoologist 11:327-339.
Packard, A. S. 1870. New or rare American Neuroptera, Thysanoptera and
Myriapoda. Proc. Boston Soc. Nat. Hist. 13:405-409.

ATURE-(Prepublished Abstract.) The effect of high temperature on
chiasma formation during oogenesis has been studied in the grasshopper
Melanoplus femur-rubrum (De Geer). Prolonged heat treatment (400 C.)
during mid-prophase of meiosis causes a reduction in the mean chiasma
frequency per cell. Only those bivalents in which more than 1 chiasma
occurs are affected by the heat. The pattern of chiasma frequency re-
sponse to heat is similar to that which occurs in males of the same species.
Heredity, 1974, 32(2): 159-164; K. Church, Arizona S. Univ., Tempe 85281.

The Florida Entomologist

MENOPTERA: FORMICIDAE)'-(Note.) Antioxidants were evaluated for
their potential for retarding the development of rancidity, a major factor affecting
field life of fire ant bait. In a modification of the method of Lofgren et al. (1961.
Imported fire ant toxic bait studies: the evaluation of various food materials. J.
Econ. Ent. 45: 1096-1100), fire ants were offered blotting paper saturated with soy-
bean oil (SBO) with and without antioxidants in replicated feeding tests. Concentra-
tions of antioxidants in the SBO ranged from 2.0 to 0.25% depending on acceptance.
Antioxidants that were unacceptable to the ants (50% reduction in feeding) included:
p-phenylenediamines (N,N'-dicyclohexyl-, N-octyl-N'-phenyl-, and N,N'-bis(1-
ethyl-3-methylpentyl)-); N-nitrosodiphenylamine; 1,2-dihydro-6-ethoxy-2,2,4-tri-
methylquinoline; polymerized trimethylquinoline; 4'-hydroxydodecananilide; 4,4'-
(2,3-dimethyltetramethylene)dipyrocatechol; p-(benzyloxy)phenol; 3,4-(methylenedi-
oxy)phenol; 2,4,6-tris[(dimethylamino) methyl]phenyl; a-(dimethylamino)cresol; CAO-
42("alkylated cresols"); and dimethyl, diethyl, or dibutyl phosphites.
The rest of the antioxidants were subjected to an accelerated oxidation treatment
involving exposure in SBO to high intensity UV (545 microwatts/cm2 of 320-400 mp at
45.6 cm) and high temperature (880C). Formulations with a low peroxide value (Anon.
1967. Active oxygen method for comparing fat and oil stabilities. Tech. Data Sheet
G-159, Eastman Chem. Prod., Inc., Kingsport, Tenn.) were then tested for acceptability
to the ants. Four hundred g lots of each formulation were exposed on sand for 24, 48, and
72 hrs to natural weathering, after which 50 g samples were offered to field colonies (5
replicates). The formulations all contained Calco@N-1700 red dye in lieu of mirex, and
acceptability was measured as the percentage of ants containing dye after 24 hrs.
Antioxidants acceptable to the ants but with high peroxide value after the accel-
erated oxidation test included: propyl gallate; butylated hydroxyanisole (butylmeth-
oxyphenols); 6-tert-butyl-2,4-xylenol; 2,6-di-tert-butyl-p-cresol; 2,6-di-tert-butyl-
a-(dimethylamino)-p-cresol; 4,4'-methylenebis[2,6-di-tert-butylphenol]; 2,2'-methylene-
bis[6-tert-butyl-p-cresol]; 4,4'-thiobis[6-tert-butyl-o-cresol]; 4,6-dinonyl-o-cresol; a,-
a',a"-(2,4,6-tetramethyl-s-phenyl)tris[1,6-di-tert-butyl-p-cresol]; dioctadecyl 3,3'-thio-
dipropionate (secondary antioxidant and synergist); triisooctyl phosphite; Mark 1089
("a substituted thiobisphenol C1, alkyl phosphite"); Hallcolife Ultra ("a hindered
phenol based on wood rosin"); and various proprietary formulations of commercial
antioxidant or other protective agents (Endox-0 ; Tenox S-10 and Tenox 7 ;
Topanol AN, Irganox 858(; Wytox 345f and Wytox 450 ).
Antioxidants not acceptable even though the peroxide value was low included
N,N-dioctyl-p-phenylenediamine; cadmium diamyldithlocarbamate; zinc bis(di-
phenyldithiocarbamate); zinc dibutyldithiocarbamate; Plastinox 1161 ("a hin-
dered phenol").
Only mono-tert-butylhydroquinone warranted testing in bait formulations. None
of the formulations with this material was superior to the controls in field tests,
but mono-tert-butylhydroquinone may provide protection against rancidity if used
in alternative formulations (alternative coatings and/or in conjunction with UV
light absorbing compounds). Further evaluation of antioxidants has been suspended
indefinitely due to priority of other research problems. The sources of these anti-
oxidants and more detailed results of the tests are available from the authors D. P.
Jouvenaz, W. A. Banks, C. S. Lofgren, and D. M. Hicks, Insects Affecting Man Re-
search Laboratory, Agricultural Research Service, USDA, Gainesville, Florida 32604.


Vol. 57, No. 3, 1974

The Florida Entomologist




Ion induced X-ray fluorescence indicated that relative total body concen-
trations of P, S, Cl, K, Ca, Ni, Cr, Ti, Mn, Fe, Cu, Zn, Pb, Sr, Mo, and Rb could
be detected in adult and immature stages of the red imported fire ant,
Solenopsis invicta Buren. In general, adult imported fire ants contained a
higher total body concentration of major and trace elements than the imma-
ture stages, with the following trend being noted: workers > pupae > larvae.

Research on the concentration and distribution of major and trace
elements in insects has been hampered by lack of suitable analytical tech-
niques having adequate sensitivity, selectivity, and practical applicability to
allow precise microanalyses of the elements accumulated in insect tissues.
Recently, techniques such as atomic absorption spectroscopy, neutron ac-
tivation analyses, and X-ray fluorescence spectroscopy have been used to
determine the concentration of various major and trace elements in insect
tissues. Levy and Cromroy (1973) have used atomic absorption spectroscopy
to determine the total body concentration of several major and trace elements
in 41 species of adult and immature insects. Ion-induced X-ray fluorescence
has been used to study the relative concentration of 14 major and trace
elements present in the tissues of 9 species of adult and immature insects (Van
Rinsvelt et al. 1973).
The purpose of this study was to compare the relative concentration of
several major and trace elements accumulated in the tissues of adult and
immature stages of the red imported fire ant, Solenopsis invicta Buren.

Two-3g samples of larvae, pupae, and a mixed sample of major and minor
workers were obtained from a red imported fire ant colony maintained at the
Insects Affecting Man Research Laboratory, ARS-USDA, Gainesville,
Tissue preparation and analyses
Each fire ant caste was freeze-killed by lyophilization and ashed in a low
temperature radio frequency furnace. A few milligrams of the ash were glued
to self-supported pure carbon films with a drop of diluted polystyrene glue.
The dried samples were then bombarded with 4 MeV helium ions produced by

Florida Agricultural Experiment Station Journal Series No. 5342.
Department of Entomology & Nematology, University of Florida, Gainesville, Florida 32611.
'Department of Physics and Astronomy, University of Florida, Gainesville, Florida 32611.

Vol. 57, No. 3, 1974

The Florida Entomologist

the University of Florida Van de Graaff accelerator and the emitted X-rays
were detected by a Si(Li) detector, and stored in a 512 channel pulse height
analyzer which finally delivered a complete spectrum of the emitted X-rays.
This permitted the identification of the elements present in the sample via
their characteristic X-rays. Specific instrumentation, techniques, and
methods of analysis were reported by Van Rinsvelt et al. (in press).

Results from ion induced X-ray fluorescence indicated that relative total
body concentrations of P, S, Cl, K, Ca, Ni, Cr, Ti, Mn, Fe, Cu, Zn, Pb, Sr, Mo,
and Rb could be detected in the adult and immature stages of S. invicta (Fig.
1-3). However, Mo and Sr in S. invicta larvae and pupae respectively could not
be distinguished from background levels and were subsequently assumed to be
present in fire ant tissues in extremely low concentrations. Levy and Cromroy
(1973) have determined the parts per million total body Cu, Fe, Na, Mg, K, and
Na in the tissues of a mixed sample of major and minor red imported fire ant
Results from pulse-height comparisons (Fig. 1-3) indicated that P, S, Cl, K,
Ti, Cr, Fe, Ni, Cu, Zn, Pb, Rb, Sr, and Mo were more concentrated in the pupal
stage than in the larval stage. However, the total body concentration of Ca in
the pupal stage was found to be approximately equal to or less than the Ca
level found in larvae. Mn was approximately equal to or greater than the level
found in larvae. All major and trace element concentrations were greater in
workers than pupae and greater in workers than larvae.
In general, adult red imported fire ants contained a higher total body
concentration of major and trace elements than the immature stages, with the
following trend being noted: workers > pupae > larvae. Results indicated that
many biologically active major and trace elements as well as environmental
contaminants can easily be detected from the ash of red imported fire ants
with the ion induced X-ray fluorescence technique. Therefore, imported fire
ant workers may serve as an effective biological indicator of metallic pollution
(e.g. Ti, Pb, Sr) in the environment.

The authors wish to thank D. P. Jouvenaz of The Insects Affecting Man
Laboratory, ARS-USDA, Gainesville, Florida for supplying the fire ants used
in the analyses. This paper was partly supported by Cooperative Agreement
Grant No. 12-14-100-10, 951 (33) entitled toxicants for control of imported fire


Levy, R., and H. L. Cromroy. 1973. The concentration of some major and trace
elements in forty-one species of adult and immature insects determined
by atomic absorption spectroscopy. Ann. Ent. Soc. Amer. 66:523-26.
Van Rinsvelt, H. A., R. Duerker, Jr., R. Levy, and H. L. Cromroy. 1973. Major
and trace element detection in insects by ion induced X-ray fluores-
cence. Fla. Ent. 56:186-90.

Vol. 57, No. 3, 1974


Levy et al.: Element Concentration in Fire Ants 271



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The Florida Entomologist

Vol. 57, No. 3, 1974







44) 0
b C




Levy et al.: Element Concentration in Fire Ants 273

A 0 IN
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The Florida Entomologist

te.) Control of the sweet potato weevil, Cylas formicarius elegantulus
(Summers), is a problem throughout the sweet potato growing areas of the
Americas. It is at times so serious that it destroys crops, making life more
difficult for those who are so dependent on sweet potatoes for food. Migration
of the hundreds of thousands of Cubans to Florida (since Castro took over
Cuba) brought about greatly increased production of sweet potatoes and of
sweet potato weevils. Fortunately dieldrin was approved for use in control, the
use of which gave measures of control.
The many hundreds of acres set with sweet potato plants helped increase
the weevil abundances. Year around production further aided weevil abun-
dance. Ipomoea sp., such as morning glory, are host plants which provide
ample infestation sources; hence, sweet potato weevil populations are very
high in Florida.
An effective, facile control method, without residue and environmental
problems, is greatly needed for the protection of sweet potatoes. Such a
method appeared available through the use of soil insecticides, those materials
that might be applied to the soil surface and control the soil inhabiting
A field experiment was started using granular materials often used to
control other soil inhabiting insects and at rates usually employed. Weevil
punctures of roots were initially counted to measure treatment results, but
later roots were put into containers and number of weevils emerging per unit
weight of the roots was determined. Insignificant and erratic results were
obtained in test after test. Other materials, increased amounts of materials,
and increased frequency of application were tried. In one test endosulfan, for
example, was most effective, in another parathion was the most effective. As
much as 1.47 kg/ha per month were tried, which with 8 applications per season
added to 11.76 kg/ha. After years of experimentation with the granular
materials and generally unsatisfactory results it was concluded that they were
not the materials to use. It is believed that the high alkalinity of the oolithic
limestone soils having pH of 7.8 to 8.3 were degrading the insecticidal
materials. Such degradations must have occurred with sufficient rapidity to
have permitted weevil infestations to develop in the vines and roots.
Two final tests were conducted in which regularly used rates of materials
were applied weekly with water. The results, although not tabulated here,
suggested that experiments along this line were needed and might be effective.
Residues may be lower from more frequent and lower rates of application,
thus tending to reduce the environmental contamination.-D. 0. Wolfen-
barger, and S. D. Walker, Agricultural Research and Education Center,
Homestead, Fla. 33030.


Vol. 57, No. 3, 1974

The Florida Entomologist



University of Florida, Agricultura-Research Center,
Fort Lauderdale 3/314

Insecticides were evaluated on tropical sod webworms, Herpetogramma
phaeopteralis Guenee, on Bermudagrass and on southern chinch bugs, Blissus
insularis Barber, on St. Augustinegrass. Carbofuran; dialifor, DyfonateT",
0-ethyl S-phenyl ethylphosphonodithioate; leptophos; and PrimicidTM, 2-
Diethylamino-6-ethylprimidin-4-yl diethyl phosphorothionate, provided ex-
cellent control of both the target webworm population and the reinfestive
population 3-4 weeks later. BiotrolT" and DipelT preparations of Bacillus
thuringiensis gave significantly better webworm control than untreated check
plots. Southern chinch bugs were controlled with treatments of carbofuran;
Dyfonate; MocapTM; 0-Ethyl S,S-dipropyl phosphorodithioate; PP484, di-
ethyl-2-ethylacetamido-6-methyl pyrimidin-4-yl phosphorothionate;
Primicid T"2-diethylamino-6-ethylprimidin-4-yl diethyl phosphorothionate;
and propyl thirpyrophosphate.

The tropical sod webworm, Herpetogramma phaeopteralis Guenee, and
the southern chinch bug, Blissus insularis Barber, are 2 of the most destruc-
tive pests of turfgrass in Florida. If left uncontrolled the chinch bug is devas-
tating to St. Augustinegrass. Webworms feed on all the southern turfgrasses,
especially on Bermudagrass and St. Augustinegrass, and can completely
remove all the green tissue if not controlled. Many insecticides (Reinert 1972a,
1972b, 1973, Stringfellow 1968, 1969) have been evaluated against these pests
but new materials which are effective need to be found. It is the purpose of this
paper to report the effectiveness of several insecticides against these 2 pests of
WEBWORMS-Infested areas of Bermudagrass were divided into 50 ft2 plots
and webworm populations sampled. A 1.5 ft buffer zone was left between plots.
Counts were made by sprinkling a quart of 0.02-0.05% pyrethrin on 4 ft2 of turf
in each plot and counting the number of worms which were driven to the
surface in 10 min. Successive samples within a plot were taken in a different
area to avoid any interaction due to the pyrethrin.
In each test the plots were divided into 4 blocks based on pre-treatment
counts and treatments were randomized within each block. Formulations and
rates for each insecticide are given in Tables 1-3. Granular materials were
dispersed with a handshaker and washed into the thatch with 2 gal of wa-
ter/plot. All other formulations were mixed with 0.5 gal of water (10 gal/1000
ft2), except Bacillus thuringienses preparations as given in Table 3, and
applied with a 2 gal compressed air sprayer.

Florida Agricultural Experimentation Station Journal Series No. 5340.

Vol. 57, No. 3, 1974


The Florida Entomologist


Larvae/4 ft2
Means* post treatment
l AI/A Pre-
(Lb) Treatmt. 4 da 1 wk 2 wk 3 wk 4 wk

Primicid 1.9G 1 70.8 1.0a 1.3 0.3 3.5a 9.6ab
Primicid 5EC 2 73.3 0.Oa 0.0 0.0 17.0ab 6.7ab
Primicid 5EC 1 73.3 0.Oa 0.0 0.0 28.3ab 16.6b
Dyfonate 2G 4 73.0 1.0a 0.0 0.0 11.8ab 12.0ab
Dyfonate 2G 2 70.8 2.0a 0.3 0.0 20.5ab 0.Oa
Dialifor 4EC 2 72.3 0.Oa 0.0 0.5 26.5ab 3.0a
Carbofuran 4F 15 72.8 0.Oa 0.0 0.5 47.0bc 12.3ab
Propyl thiopyrophosphate 6E 3.8 71.3 6.5b 1.3 0.0 65.3cd -
Methomyl 90 WP 1 70.8 1.0a 0.0 2.8 68.0cd
Methomyl 90 WP 2 75.5 0.5a 0.0 2.5 111.3d
Methomyl 1.8L 1 74.0 0.Oa 0.0 0.3 109.0d -
Methomyl 1.8L 2 71.5 1.8a 0.0 0.0 85.3d
Largon 25W 0.3 76.0 2.5b 0.0 0.5 77.0d
Gardona 75WP 1 72.5 3.0b 0.5 0.0 90.0d -
Phenthoate 4E 5 73.5 2.8b 2.0 0.8 97.2d -
Chlorpyrifos 2E 1 74.5 1.5a 0.3 0.5 134.5d
Untreated check 0 72.3 19.3c 12.8 2.0 78.0d 26.0c

*Means in a column not followed by the same letter are significantly different (P =.05) (Duncan's
multiple range test).
CHINCH BUGS-Treatments were applied to 100 ft2 plots of St. Augus-
tinegrass showing chinch bug damage. Populations were sampled before in-
secticides were applied by forcing a metal cylinder equivalent to 1 ft2 into the
turf, filling it with water, and counting the bugs that surfaced in 8 min. Plots
were divided into blocks and treatments randomized as above. Granular
materials were dispersed with a hand shaker and washed into the thatch with
6 gal of water/plot. Other formulations were mixed with 2 gal of water, applied
with a sprinkler can, and washed in with 4 gal of water/plot. Formulations and
rates are presented in Tables 4-5.
INSECTICIDEs-Chemical definitions of the proprietary compounds used in
the tests are:
BiotrolTM, Bacillus thuringiensis Berliner
DipelTM, Bacillus thuringiensis Berliner
DyfonateT", 0-ethyl S-phenyl ethylphosphonodithioate
GardonaT, 2-Chloro-l-(2,4,5-trichlorophenyl vinyl dimethyl phos-
ImidanTM, N-(Mercaptomethyl) phthalimide S-(0-0-dimethylphos-
MocapTM, 0-Ethyl S,S-dipropyl phosphorodithioate
PrimicidTM, 2-Diethylamino-6-ethylprimidin-4-yl diethyl phosphor-
PP-484, diethyl-2-ethylacetamido-6-methyl pyrimidin-4-yl phos-
LargonT 1-(4-chlorophenyl)-3-(2,6-difluorobenzoyl)-urea

Vol. 57, No. 3, 1974


Reinert: Sod Webworm and Chinch Bug Control



Larvae/4 ft'

Means* post treatment

AI/A Pre-
Chemical (Lb) Treatmt. 2 da 1 wk 2 wk 3 wk 4 wk 5 wk

Leptophos 2.7EC 1 108.0 0.5a 0.0 0.0 0.0 0.Oa 0.0
Primicid 5EC 2 100.5 3.0ab 0.3 0.0 0.0 0.5a 0.3
Methomyl 90WP 2 109.2 1.3a 0.5 0.0 1.8 5.8bc 6.5
Mocap 69EC 1 103.0 18.5c 6.5 1.3 1.8 1.5a 3.8
Mocap 69EC 2 102.0 11.8c 5.3 0.3 2.5 5.0bc 3.3
Mocap 10G 2 106.5 14.3c 4.5 0.5 0.0 2.8ab 2.5
Dimethoate 4EC 1 104.2 36.5d 18.0 1.0 3.5 1.8a 1.8
Imidan 50WP 2 106.5 48.5d 15.2 0.3 1.3 3.5ab 2.0
Gardona 75WP 1 106.0 18.5c 3.3 0.0 0.3 7.0bc 6.8
Acephate 75WP 1 99.3 7.3bc 7.0 0.0 2.5 7.0bc 1.8
Untreated Check 0 106.7 36.0d 8.0 3.8 6.8 8.5c 3.3

*Means in a column not followed by the same letter are significantly different (P =.05) (Duncan's
multiple range test).


Larvae/4 Ft'


Material/A Spray Pre-
(Lb) 1,000 ft Treatmt.

Chlorpyrifos 2E
Dipel WP
Dipel WP
Dipel WP
Dipel WP +
Corn Oil**
Dipel WP
Dipel WP
Biotrol XK WP
Biotrol XK WP
Biotrol Bait
Untreated Check

Means post treatment

3 da wk 2wk 3wk



*Grass recovery ratings scaled 1-10; where 1 = no recovery, and 10 = complete recovery. Means in
a column not followed by the same letter are significantly different (P=.05) (Duncan's multiple
range test).
**Corn oil added at 1 quart/acre.
tLbs AI/A.



4 wk 1 wk





The Florida Entomologist

Vol. 57, No. 3, 1974

ON 25 MAY 1973 (4 REPLICATES).

Chinch bug numbers/Ft2

AI/A Pre-
(Lb) Treatmt.

Means** at weeks post treatment

2 3 4 5 6

phosphate 1.1EC 3.7 88.8 2.0 0.3 0.5 0.3a 1.5 1.0a
phosphate 1.1EC 7.3 95.3 0 0.3 2.5 0.8a 2.8 1.3a
Dyfonate 2G 4 87.5 0.8 0 0.5 0.5a 1.0 5.0a
Carbofuran 10G 10 97.0 7.0 0.8 0.5 0.8a 1.5 1.5a
PP-484 2.1EC 2 88.8 9.8 2.3 8.8 4.0a 5.5 2.5a
Primicid 5EC 2 87.5 0.3 0 0.5 1.0a 2.5 4.8a
Primicid 5EC 1 91.2 36.4 16.8 29.8 17.2b 29.6 25.3b
Untreated Check 0 90.0 82.0 82.0 78.3 85.0c 94.0 79.3c

*Populations in excess of 100 chinch bugs were recorded as 100.
**Means in a column not followed by the same letter are significantly different (P=.05) (Duncan's
multiple range test).



Chinch bug numbers/Ft2

Means** at weeks post treatment

AI/A Pre-
Chemical (Lb) Treatmt. 1 2 3 4 5 6

Mocap 10G 10 93.0 7.8 1.3 9.8 9.8a 32.0 36.3a
Mocap 6E 10 90.0 1.8 3.5 15.8 17.5ab 11.8 12.8a
Dimethoate 2E 2.5 90.0 28.5 25.5 23.5 22.0ab 25.8 29.8a
ride 95WP 1 81.5 34.8 59.5 50.0 46.5b 44.5 17.8a
Phenthoate 4E 2.5 88.5 21.3 44.5 50.0 51.5b 54.8 51.5a
Untreated Check 0 100.0 87.8 87.8 100.0 100.Oc 100.0 100.0b

*Populations greater than 100 were recorded as 100.
**Means not followed by the same letter are significantly different (P=.05) (Duncan's multiple
range test).




Reinert: Sod Webworm and Chinch Bug Control 279

WEBWORMS-A comparison of control by 10 insecticides is given in Table 1.
Very good control was provided by all the chemicals 4 days after treatment. A
comparison was also made between different rates and formulations of
Dyfonate, methomyl, and Primicid. It appears that the granular formulation
of Primicid was more effective than the emulsifiable concentrate. Both rates
of Dyfonate were equally effective. Methomyl was equally effective for both
rates and formulations. Only treatments of carbofuran, dialifor, Dyfonate,
and Primicid were significantly better than the untreated check at 3 weeks
when reinfestation occurred. Only plots treated with these 4 chemicals
remained green at 4 weeks. All other plots were completely stripped by the
high density of second generation larvae.
In test 2 (Table 2) 8 chemicals were compared for control of webworms. All
the materials gave control by 1 week. Leptophos gave good residual control as
did the 4 compounds in the above test. Plots were reinfested by much smaller
populations than in the previous test.
In test 3 (Table 3) a comparison was made of Dipel and Biotrol at different
gallonage applications and rates of applications. Only minor differences were
apparent due to different rates of water. Both materials gave significantly
better control than the untreated check when the plots were visually rated for
recovery at 1 week. Bacillus treatments were not as good as the treated check
plots of chlorpyrifos.
In conclusion, it is apparent that most of the materials used gave good
control of the target generation of worms, but only carbofuran, dialifor,
Dyfonate, leptophos, and Primicid gave residual control of the second
generation population.
CHINCH BUGS-Results for 9 insecticides evaluated for southern chinch bug
control are given in Tables 4-5. Control was provided by applications of
carbofuran, Dyfonate, Mocap, PP-484, Primicid, and propyl thiopyro-
phosphate. From Table 4 it appears that the half rate of propyl thiopyro-
phosphate will give good control. The higher rate of Primicid is necessary
for this pest.
These tests were designed to allow reinfestation of the treated plots as soon
as possible after treatment to determine the maximum residual activity for
each insecticide. Materials which provided control in these tests would
probably provide much longer control under field conditions where the entire
infested area was treated.


Reinert, J. A. 1972a. Control of the southern chinch bug, Blissus insularis, in
South Florida. Fla. Ent. 55:231-5.
Reinert, J. A. 1972b. Turf-grass insect research. Proc. Fla. Turg-Grass Manag.
Conf. 20:79-84.
Reinert, J. A. 1973. Sod webworm control in Florida turfgrass. Fla. Ent.
Stringfellow, T. L. 1968. Studies on turfgrass insect control in South Florida.
Proc. Fla. State Hort. Soc. 81:447-54.
Stringfellow, T. L. 1969. Developments in Florida turfgrass insect control,
1969. Proc. Fla. Turf-Grass Manag. Conf. 17:94-100.

The Florida Entomologist

SCALES'-(Note.) When handling quantities of Anticarsia gemmatalis
Hiibner, I found the tufts of scales on the male legs very obvious in distin-
guishing male moths from females at a distance. The long scales (Fig. 1) were
present on the femur of the prothoracic legs and tibia of the metathoracic legs
of the male. Scales were short and sparse on the female legs.
Since there was variation in the presence of leg scales, 200 moths were
separated according to the presence or absence of the leg scales. In checking
the genitalia of these moths, 99% had been correctly determined with respect
to gender. The 1% incorrectly determined moths were males which had lost
most of their body scales.
Several thousand moths from field collections and laboratory colonies
have been sexed by the leg scales. These moths vary considerably in wing color
from light tan to almost black. Groups of moths from a field location or a
laboratory reared group tended to segregate, with the lighter tan moths being
females and the darker moths being males. However, this relative character is
not as dependable or accurate a means of determining sex as is the leg scale

'Florida Agricultural Experiment Station Journal Series No. 5408.




Fig. 1.-Scale tufts on the meso- and pro-thoracic legs of the male and absence on the female legs.


Vol. 57, No. 3, 1974

Vol. 57, No. 3, 1974 The Florida Entomologist 281




Populations of Plecia nearctica Hardy were surveyed during the spring
and fall of 1973 to monitor adult flights which usually occur in May and
September. The May adult flight was noticeably smaller. A large flight was
observed in September. During early spring, 5 species of fungi were isolated
and identified from dead and moribund lovebug larvae collected in the
Gainesville area. In the fall, 2 additional fungal species were isolated, 1 from
adults during the September flight and 1 from laboratory reared larvae.

Each year, large numbers of adult lovebugs, Plecia nearctica Hardy,
emerge from grass litter in central and north Florida. According to Hetrick
(1970), the adults emerge in 2 yearly flights of about 4 weeks duration in May
and September. They are a safety hazard to drivers as dead adult insects may
cover auto windshields, impairing driver vision.
In the spring of 1973, there was a noticeable decrease in the lovebug larva
population, resulting in a reduced adult flight in May. Field observations
demonstrated a high percentage of late larval instars apparently killed by
fungi. The usually dense covering of bright colored, fungal hyphae made their
mummified bodies readily observable. A survey was conducted to determine
the identity of the fungi involved.

Field collections in pastures and on road banks were made at weekly
intervals from April to September. Collection sites were selected based on
good drainage and proximity to lakes, ditches, and roadside pools. Dead and
moribund larvae were collected and transported to the laboratory for
examination. Cadavers were placed in a moist chamber and held at 260 C until
fungal sporulation occurred. No surface sterilization was used.

Five species of fungi were routinely isolated from larvae collected during
April and May. These isolates included Beauveria bassiana (Bals.) Vuill.,
Metarrhizium anisopliae (Metsch.) Sorok., Eupenicillium brefeldianum
(Dodge) Stolk & Scott, Tolypocladium cylindrosporum W. Gams, and
Apiosordaria verruculosa (Jensen) von Arx and W. Gams. The 2 fungi
isolated in the fall included Conidiobolus coronatus (Cost.) Batko identified
from adults during the September flight, and Arthrobotrys oligospora Fres.,
from field collected, laboratory reared larvae.

Florida Agricultural Experiment Station Journal Series No. 5263.
SDepartment of Entomology and Nematology, University of Florida, Gainesville, Florida 32611.
SDepartment of Botany, University of Florida, Gainesville, Florida 32611.

The Florida Entomologist

Vol. 57, No. 3, 1974

;"d %"iI


r I,
* ^ i^B^^^





Fig. 1. Beauveria bassiana. Conidiogenous cells with abundant conidia. x
675. Fig. 2. Metarrhizium anisopliae. Conidiophores, phialides, and conidia. x
675. Fig. 3. Conidiobolus coronatus. Spores. x 275. Fig. 4. Eupenicillium
brefeldianum. Superficial cleistothecia on agar. x 675. Fig. 5. Eupenicillium
brefeldianum. Spores. x 675. Fig. 6. Arthrobotrys oligospora. Conidiophore
showing septate conidia. x 675.
Beauveria bassiana (Fig. 1), M. anisopliae (Fig. 2), and Conidiobolus
coronatus (Fig. 3) are well known entomopathogenic fungi (Steinhaus and
Marsh 1962, Madelin 1966). All of these genera infect a wide range of insect



v I
,Ab 4

Kish et al.: Fungi Associated with Plecia nearctica

hosts (Charles 1941, Leatherdale 1958, Madelin 1966, MacLeod and Muller-
Kbgler 1973).
Eupenicillium brefeldianum (Fig. 4,5) was first described from a human
alimentary tract as Penicillium brefeldianum (Dodge 1933). After continuous
propagation in the laboratory, it rapidly lost its ability to form conidia. It now
forms abundant cleistothecia (Fig. 4), but even these abort development
before spores are formed.

at~- 4jt

2. -I- .


Fig. 7,8,9. Cladorrhinum state of.Apiosordaria verruculosa. Phialides and
spores. x 675. Fig. 10. Tolypocladium cylindrosporum. Phialide habit. x 675.

Arthrobotrys oligospora (Fig. 6) is a predaceous fungus known to attack
soil nematodes (Dreschler 1937). It is easily maintained in the laboratory, and
forms abundant conidia. An attempt was made to rear approximately 25
larvae, hatched from field collected eggs, on moist filter paper. The fungus
entangled the young larvae, usually trapping 1 or 2 at a time, and rendered
them immobile. Individual hyphae were attached firmly to the integument.
Death occurred within 3 days after entrapment. The cycle continued for 2
weeks, until all larvae succumbed.




The Florida Entomologist

Apiosordaria verruculosa is a heterothallic soil inhabitant with a
Cladorrhinum imperfect state (Fig. 7,8,9) (von Arx and Gams 1966).
Tolypocladium cylindrosporum is known only as a soil saphrophyte (Gams
1971). It is similar to the entomogenous genus Beauveria Vuill.; however,
Beauveria forms conidia sympodially, while Tolypocladium is a philalide
producer. A sister species, T. inflatum Gams, was isolated by E. Miiller-Kogler
from Aradus cinnamomeus. Although T. cylindrosporum is not known to be
pathogenic, it is the organism most frequently isolated from lovebug larvae.
Preliminary indications are that the failure of a lovebug flight to develop in
the spring of 1973 was associated with the activity of all, or some, of these
fungi. Re-infectivity studies will determine the pathogenicity of all 8 fungus
species and the parameters required for their development. Further research
will be conducted to determine the role of these fungus species as natural
control agents of the lovebug.


Charles, V. K. 1941. A preliminary check list of the entomogenous fungi of
North America. USDA Bur. Plant Ind. Inst. Pest Survey Bull.
Dodge, B. 0. 1933. The Perithecium and Ascus of Penicillium. Mycologia.
Dreschler, C. 1937. Some hyphomycetes that prey on free-living terricolous
nematodes. Mycologia. 29:447-552.
Gams, W. 1971. Tolypocladium, eine Hyphomycetengattung mit geschwollen
Phialden. Persoonia. 6:185-191.
Hetrick, L. A. 1970. Biology of the "Love-bug", Plecia nearctica (Diptera:
Bibiondae). Fla. Ent. 53:23-26.
Hoog, G. S. de. 1972. The genera Beauveria, Isaria, Tritirachium and
Acrodontium. Gen. Nov. CBS. Studies in Mycology, No. 1, 1-41.
Leatherdale, D. 1958. A host catalogue of British entomogenous fungi. Ent.
Mon. Mag. 94:103-105.
MacLeod, D. M., and E. Miiller-Kogler. 1973. Entomophthera species with
pear-shaped to almost spherical conidia (Entomophthorales: En-
tomophthoracae). Mycologia 55:823-93.
Madelin, M. F. 1966. Fungal parasites of insects. Ann. Rev. Ent. 11:423-448.
Steinhaus, E. A., and G. A. Marsh. 1962. Report of diagnoses of diseased
insects 1951-1961. Hilgardia. 33(9).


Vol. 57, No. 3, 1974

The Florida Entomologist

series of specimens of Eriococcus smithii Lobdell was collected 19 October
1973 from broom sedge, Andropogon virginicus L., at Tall Timbers Research
Station, Leon County, Florida. This series, identified by D. R. Miller,
Systematic Entomol. Lab., ARS, USDA, Washington, D. C., is the first record
of this species occurring in Florida. The species was originally described by
Lobdell (Ann. Ent. Soc. Amer., 1929, 22:764-5) from specimens collected on
broom sedge at Meridian, Mississippi by M. R. Smith. These were being tended
by the Argentine ant, Iridomyrmex humilis Mayr.
The specimens of E. smithii taken in Leon County were from 7 clumps of
broom sedge located within 5 m of each other in a shallow ditch between a
forest road and soybean field. When first collected in October 1973, these were
being tended by the dolichoderine ant Conomyrma insana (Buckley). No
other colonization of broom sedge by the scale was found in the locale. The
scale colonies on the broom sedge were intermittently checked for spread to
other broom sedge clumps and for population increases through September
1974. By September 1974, colonized broom sedge clumps had increased to 43,
spaced out over approximately 30 m within the shallow ditch. Thirty-nine of
the colonies were being tended by Conomyrma insana. Four colonized clumps
of broom sedge were located outside of the nesting area and foraging range of
C. insana. These scales were continuously tended by workers from a single
colony of the red imported fire ant, Solenopsis invicta Buren, located 13 m
from the nearest scale-colonized clump of broom sedge. J. C. Nickerson, and
W. H. Whitcomb, University of Florida, Gainesville, 32611; and G. W. Dekle,
Department of Plant Industry, Gainesville, Florida 32601.

Vol. 57, No. 3, 1974

The Florida Entomologist

TION BY RAINBOW TROUT-(Notice.) Samples of the invertebrate
drift on the surface of a eutrophic prealpine stream were collected over 6 days
in September 1970 with a floating trap. Eighty-five percent of the drift were
emerging and egg-laying aquatic insects, mainly Ephemeroptera and
Chironomidae; the remaining 15% was composed of winged terrestrial insects
like Aphidae and ants. The diel pattern of the drift showed 2 strongly marked
peaks; the first, at about 15.00 h, being caused by emerging Ephemeroptera,
the second, at about 19.00 h, by egg-laying adults of the mayfly Caenis
macrura and by Chironomidae.
The composition of the drift was compared with the composition of the
stomach contents of rainbow trout (Salmo gairdneri) of age classes 2 + to 4 +
caught during the same time period and in the same part of the stream. The
trout were exploiting only the surface insect drift.
Good correlation was found between the diel variations in the abundance
of most insect groups in the drift and in the stomachs.
The trout fed selectively within the drift: a very close correlation was
found between the mean body length of the insects and the logarithm of the
electivity coefficient (forage ratio). A further important criterion of selection
is the degree of familiarity with the prey organisms. The relative abundance of
the different prey species has no influence on the selective behaviour.
The importance of environmental factors in the choice of food sources and
for the selective feeding behaviour of trout and the possible influence of the
selective feeding on the composition of the invertebrate stream fauna are
discussed. Oecologia (Berl.) 1974, 14, 247-267; J-P. Metz, Univ. Munchen.

Vol. 57, No. 3, 1974

The Florida Entomologist



Cotton Insects Research Laboratory,
Agric. Res. Serv., USDA, Brownsville, Texas 78520


Thiotepa, a triaziridinyl phosphinic sulfide, was the most effective of 75
compounds evaluated as chemosterilants against the tobacco budworm,
Heliothis virescens (F.). Thiotepa was applied topically to adults, and it was 4
times more effective against males than females. Females treated with
thiotepa and placed in competition with untreated females were competitive
at 4, 2, and 1:1 ratios; males handled similarly were competitive at a 1:1 ratio
only. However, males treated with thiotepa and untreated males transferred
eupyrene sperm equally well to untreated females. Two heteroaromatic
aziridines, tretamine (an s-triazine) and 2,6-bis(1-aziridinyl)-pyrazine (a
pyrazine), sterilized both sexes when applied topically. The same dose of
tretamine sterilized both sexes without inhibiting sexual competitiveness.
Pairs of adults from larvae that had been injected or fed with the
nonaziridinyl methylenedioxy compound, 5-acetyl-5-[4,5-(methylenedioxy)-
2-propylbenzyl]-4,5-dihydrofur-2(3H)-anone, were sterile; pairs composed
of treated moths and untreated moths were not.

A chemical that would sterilize both sexes of the tobacco budworm,
Heliothis virescens (F.), at the same concentration without destroying the
sexual competitiveness of the insects would be useful in sterile release
Soto and Graves (1967) and Flint et al. (1968a) evaluated the sterilizing
effects of topical applications of a variety of aziridinyl and nonaziridinyl
compounds against tobacco budworms; Soto and Graves (1967), Flint et al.
(1968b), and Bankwith and Graves (1970) tested the effect of feeding several
compounds to adults of this species in the laboratory. All references indicated
that 1 or more aziridinyl compounds were effective.
Bowers (1968) showed that some methylenedioxyphenyl (MDP) com-
pounds possessed a high order of juvenile hormone activity and were active
against certain species of insects. Guerra et al. (1973) reported that 3 juvenile
hormone analogs with the MDP moiety were effective chemosterilants when
applied topically or fed to the tobacco budworm.
In 1972-73, at the Brownsville laboratory, we evaluated the sterilizing
effects of 75 compounds applied to tobacco budworm adults or larvae as

Heliothis virescens (F.) (Lepidoptera: Noctuidae).
In cooperation with the Texas Agricultural Experiment Station, Texas A&M University,
College Station 77843.
This paper reports the results of research only. Mention of a pesticide or a proprietary product
does not constitute a recommendation or an endorsement by the USDA.
SMetabolism and Irradiation Research Laboratory, Fargo, N. D. 58102.

Vol. 57, No. 3, 1974


The Florida Entomologist

topical, injection, or diet-incorporated treatments. Included among the 75
compounds were certain aromatic aminoethanols known to be starting
materials for synthesis of the cyclic aziridine compounds (Rudesill et al. 1971).
We hypothesized they might undergo cyclization in vivo to compounds capa-
ble of sterilizing the tobacco budworm.
Our objective was to determine the sterilizing ability of each compound
within the limits of its solubility. Our criterion for sterility was 90% or more
reduction in egg hatch from that of the untreated check. We also determined
competitiveness of moths treated with thiotepa and tretamine.

Tobacco budworms used in these experiments had been reared in our
laboratory for at least 3 years prior to these experiments.
All tests described herein were held at 27+20 C, 65+5% RH, and natural
light. Test compounds 1-12, 26-38, and 40-75 when applied topically to
adults or larvae were diluted in acetone. Compound 39 was diluted in
methanol+water (with 1% Triton X-155) in a 1:1 ratio. The chemical
names of the test compounds (asterisk indicates previously tested against
this insect by other investigators) are as follows:
*1. P,P-bis(l-aziridinyl)-N-methylphosphinic amide
2. P,P-bis(l-aziridinyl)-N,N-dimethylphosphinic amide
3. P,P-bis(l-aziridinyl)-N-ethylphosphinic amide
4. P,P-bis(l-aziridinyl)-N-propylphosphinic amide
5. P,P-bis(l-aziridinyl)-N-isopropylphosphinic amide
6. P,P-bis(l-aziridinyl)-N-butylphosphinic amide
7. P,P-bis(l-aziridinyl)-N-octylphosphinic amide
8. P,P-bis( -aziridinyl)-N-[4,6-bis(dimethyl amino)s-triazin-2-
yl]phosphinic amide
*9. tepa
10. thiotepa
*11. metepa
*12. hempa
13. 2-(p-nitroanilino)ethanol
14. N-(2-bromoethyl)-p-toluidine monohydrobromide
15. 2-(p-nitroanilino)ethanolp-toluenesulfonate
16. 2-(p-chloroanilino)ethanol
17. N-(2-bromoethyl)-p-fluoroaniline monohydrobromide
18. p-[(2-bromoethyl)amino]phenol
19. N-(2-chloroethyl)-p-anisidine
20. 2-(p-chloroanilino)ethanol p-toluenesulfonate
21. N-(2-bromoethyl)aniline
22. N-(2-bromoethyl)-p-anisidine
23. N-(2-bromoethyl)-p-chloroaniline
24. N-(2-bromoethyl)-p-toluidine monohydrobromide
25. p-bromo-N-(2-bromoethyl)aniline
26. l-(p-nitrophenyl)aziridine
27. l-(p-methoxyphenyl)aziridine
28. p-(l-aziridinyl)benzonitrile
29. 2,6-bis(l-aziridinyl)pyridine
30. 1-(p-fluorophenyl)aziridine
31. 1-p-tolylaziridine


Vol. 57, No. 3, 1974

Wolfenbarger et al.: Sterilants for H. virescens

32. 1-phenylaziridine
33. 4'-(l-aziridinyl)acetophenone
*34. 2,6-bis(l-aziridinyl)-pyrazine
35. 4-chloro-2,6-bis(dimethylamino)pyrimidine
36. 2,4,6-tris-aziridinyl-pyrimidine
37. 4-(l-aziridinyl)-1,2-naphthoquinone
38. hemel
*39. tretamine
40. 1,2-(methylenedioxy)benzene
41. 3,4-(methylenedioxy)chalcone
42. l-bromo-3,4-(methylenedioxy)benzene
43. 3,4-(methylenedioxy)-l-nitrobenzene
44. [3,4-(methylenedioxy)phenyl]acetic acid
45. 3,4-(methylenedioxy)-/f-nitrostyrene
46. 3,4-(methylenedioxy)benzonitrile
47. 1-[3,4-(methylenedioxy)phenyl]-2-nitro-l-propene
48. 3',4'-(methylenedioxy)acetophenone
49. 3,4-(methylenedioxy)aniline HCL
50. 3,4-(methylenedioxy)benzylamine
51. N-[3,4-(methylenedioxy)benzylidene]aniline
52. 3,4-(methylenedioxy)cinnamic acid
53. 3,4-(methylenedioxy)mandelic acid
54. 3,4-(methylenedioxy)phenol
55. piperonyl butoxide
56. propyl some
57. sesamex
58. sulfoxide
59. 4,5-(methylenedioxy)-2-propylbenzyl laurate
60. 5-acetyl-5-[4,5-(methylenedioxy)-2-propylbenzyl]-4,5-
61. a-(ethylsulfonyl)-4,5-(methylenedioxy)-2-propyltoluene
62. 4,5-(methylenedioxy)-2-propylbenzyl 2-benzoylbenzoate
63. 4,5-(methylenedioxy)-2-propylbenzyl 4-ethoxybenzoate
64. 5-methyl-4-[3,4-(methylenedioxy)phenyl]-1,3-dioxane
65. 1,2-(methylenedioxy)-4-propyl-5-(2-propynyloxy)benzene
66. 7,11-dimethyl-1-[3,4-(methylenedioxy)phenyl]-1,4,6,10-
67. 3-methyl-6,7-(methylenedioxy)-1,2,3,4-tetrahydronaphth-
68. N-[2-acetoxy-3-[3,4-(methylenedioxy)phenyl]propyl]acetamide
69. cis-5-[(4,5-(methylenedioxy)-2-propylbenzyl)oxy]-2-phenyl-
70. piperonal bis[2-(2-butoxyethoxy)ethyl] acetal
71. 4-chloro-2-nitrophenyl 2-propynyl ether
72. 2-propynl 2,4,5-trichlorophenyl ether
73. 2-nitrophenyl 2-propynyl ether
74. N-(2-ethylhexyl)-5-norbornene-2,3-dicarboximide
75. bis(2,3,3,3-tetrachloropropyl) ether
When adults were treated, compounds 1-12 and 26-39 were applied
topically with a commercial automated microapplicator to the dorsum of the


The Florida Entomologist

abdomen; 8-16 male or female (1 day old), reared from laboratory stock, were
treated with each dose and crossed with untreated moths of the opposite sex.
Each dose was replicated 4-8 times, thus 2 treated males or females were
paired with 2 untreated moths of the opposite sex; also, 2 pairs of untreated
moths were used as checks for each chemical. The groups of 4 moths were
placed in separate (ca. 1-liter) cylindrical cardboard containers and fed 10%
sugar solution. The cheesecloth pads used to cover the containers served as
oviposition sites and were replaced daily during the life of the moths. A portion
of the eggs was counted and held to determine percentage hatch. Mating was
determined by the presence of spermatophores in the bursa copulatrix of the
female (Wolfenbarger and Guerra 1971).
Larvae weighing 180-200 mg were treated topically with compounds 40-42,
44, 50, and 55-75 on the dorsum of the thorax or they were injected ventrally
with compounds 40-43, 45-47, 50, 55-75 at the junction between the me-
tathorax and 1st abdominal segment (Wolfenbarger and Lowry 1969). Larvae
were treated with 100 pg of compound dissolved in 1 [1 of acetone.
Test compounds 13-25 and 40-75 were incorporated with a liquifier blender
into standard larval diet to give an 0.25% (wt/vol) concentration (Guerra
1970). Ca. 15 ml of each diet were poured into 3/4-oz transparent plastic cups
and infested the next day with one Ist-instar larva each.
Single pairs of adults from all treatments were placed (5-20 pairs/treat-
ment per chemical) in ca. 1-liter wide-mouth jars, provided with cheesecloth
covers and resting sites, and fed 5% sucrose solution (Guerra 1970).
Checks consisted of 5 or 6 couples of untreated moths for each treatment.
Each cross was replicated 4 times. Fecundity and fertility were measured and
recorded as described by Guerra et al. (1973).
We tested the sexual competitiveness of moths treated with thiotepa (10)
and tretamine (39) by combining males or females in ratios of 1:1:1, 2:1:1, and
4:1:1 treated and untreated moths of the same sex to untreated moths of the
opposite sex. The number of moths per cage was constant (12 moths/3-liter
glass globe) as described by Wolfenbarger et al. (1973). The percentage of
observed egg hatch vs. expected hatch was compared visually and by X2
analysis, and the percentage of expected egg hatch was calculated as described
by Fried (1971).
Also, estimates were made of the amount of eupyrene sperm in the sper-
mathecae of untreated females and females treated with 16 [tg thiotepa. The
single-pair reciprocal crosses of 1-day-old treated and untreated moths were
arranged; checks consisted of both treated and untreated crosses. The pairs
were held in individual (ca. 0.5-liter) cardboard cartons; eggs on the
cheesecloth cover were collected and counted the 2nd through the 5th day;
hatch was recorded. When the moths were 8 days old, the spermathecae were
excised, squashed, and observed with phase contrast microscopy as described
by Flint and Kressin (1969). Moderate to large amounts of eupyrene sperm
were designated as normal, based on observations of over 1000 squashes of
spermathecae prior to this experiment. The spermatophores were counted.

Data for the compounds that were active when applied topically to adults
are reported in Table 1. Compounds 4, 6-8, 12, 26-28, 30-33, and 35-38 were


Vol. 57, No. 3, 1974


Wolfenbarger et al.: Sterilants for H. virescens 291


Egg hatch

Avg no./
Chemo- Dose %
steri- Sex (Ag/ Eggs Spermato- reduction
lant* treated moth) observed phores % from check




*See text for code of chemical compound.

Compound 5, a branch chain homolog, sterilized males but not females of
the tobacco budworm, and it was a more effective chemosterilant than its
straight chain homolog, compound 4 (Table 1). Compounds 1, 2, and 3 showed
some activity. Compounds 9, 11 (metepa), and 29 (a pyridine) sterilized males
but not females. Only compounds 10 (thiotepa), 34 (a pyrazine), and 39 (tre-
tamine) sterilized both sexes and were not toxic to the moths nor did they

The Florida Entomologist

affect mating frequency (spermatophores/females), fecundity, or longevity
(data not shown in table); only tretamine sterilized both sexes at the same
The rationale for the evaluation of the aromatic aminoethanols (com-
pounds 13-25) was either incorrect or the cyclic end products failed to reach a
critical site.
Pairs of moths injected as larvae with compounds 60-63 and 67 failed to
reproduce (check moths had 70% hatch). Pairs treated topically as larvae with
compounds 59, 64, and 67 had 97-100% less egg hatch than the check (check
moths had 60% hatch). Pairs treated by feeding compound 60 in the larval diet
had no hatch (check moths had 71% hatch). Compound 40,
methylenedioxybenzene, the basic MDP, was inactive as a chemosterilant
when applied by any of the 3 methods used in these tests. None of the
compounds was toxic when injected or applied topically, but compounds 40,
45, 56, 58, 61, 64, 65, 67, 69, 71-73, and 75 were toxic when they were added to
the larval diet. (Because they were either toxic or ineffective as
chemosterilants when added to the diet, compounds 43 and 45-47 were tested
by injection only, compound 44 by topical application only, and compounds
48, 49, and 51-54 by neither method. Compounds 41, 42, 50, 55, 57, 66, 68, 70,
and 74 were ineffective by all 3 methods of treatment.)
Adults of both sexes from larvae reared on diets containing 0.25% (wt/vol)
of compound 60 were not sterile when they were crossed with untreated moths
of the opposite sex. There was 37% less hatch than the check (which had 54%
hatch) when treated females were crossed with untreated males. Thus, we
conclude that this compound was not effective as a chemosterilant for the
tobacco budworm at the conditions of our test.
The females treated with thiotepa (10) were competitive with untreated
females (Table 2) as indicated by X2 analysis. The observed percentage egg
hatch was lower than the expected percentage egg hatch. Guerra et al. (1972)
also found that female tobacco budworms sterilized by a combination of
reserpine and irradiation were competitive. Thus, we have 2 methods of
sterilizing female tobacco budworms that do not destroy competitiveness with
untreated females for untreated males under laboratory conditions.
The males treated with thiotepa were not competitive with untreated
males at ratios of 4 treated to 1 untreated and 2 treated to 1 untreated, but
they were competitive at a ratio of 1 treated to 1 untreated. Wolfenbarger et
al. (1973) and Wolfenbarger and Guerra (1972) obtained similar results with
tobacco budworms that had been irradiated in the egg or pupal stage, respec-
tively. We have no explanation for these phenomena.
No difference was apparent in mating frequency among the crosses or in
the sperm content of the spermathecae of treated or untreated moths after 1
or more matings (Table 3). Thus, eupyrene sperm reached the spermathecae of
at least 50% of the females in all crosses; however, the moths treated with
thiotepa were sterile.
Chi square analysis indicated that tretamine-treated males and females
were competitive with untreated males and females at all ratios (Table 4).
Moths treated with tretamine (39) (48 ptg/moth) or untreated (35 single
pairs/cross) showed little difference in niating frequency or fecundity among
the crosses; mating frequency ranged from 2.8 to 2.6 spermatophores/females
and fecundity was 454, 373, and 479 eggs/female for the treated male X
untreated female, treated female X untreated male, and untreated female X
male crosses, respectively.


Vol. 57, No. 3, 1974

Wolfenbarger et al.: Sterilants for H. virescens



No. of moths/ Avg no./ Egg hatch
Eggs Spermato- % ob- % expected
T : T : U : U 6 observed phores served ([E] x 100)**

Treated Males
8 0 2 2 114 3.3 44 14s
6 0 3 3 88 4.1 35 22s
4 0 4 4 92 2.8 39 32 ns
6 0 0 6 73 3.7 2

Treated Females
0 8 2 2 53 1.3 11 14ns
0 6 3 3 52 2.3 17 22 ns
0 4 4 4 55 2.8 25 32 ns
0 6 6 0 61 3.6 2
0 0 6 6 79 3.6 62

*T= Treated; U = Untreated.
**Computed by methods of Fried (1971); XZ was computed for percentage observed vs. percen-
tage expected hatch; s indicates significance at 1% level of probability and ns indicates no sig-
nificance at 5% level of probability.


% moths with Eggs
normal amounts
of eupyrene No.
Cross No. of sperm in spermato- Avg
( x 9 ) pairs spermatheca* phores/ 9 no./ 9 % hatch




*Following description by Flint and Kressen (1969) as "normal complement" and based on over 1,000
squashes of spermathecae prior to this experiment.

Although ca. 70% as many females of treated crosses as check females
received normal complements of eupyrene sperm, egg hatch was only 0.1% in


The Florida Entomologist



No. of moths/ Avg no./ Y Egg hatch
Eggs Spermato- % ob- % expected
Td : T : U : U9 observed phores served ([E] x 100)**

Treated Males
8 0 2 2 107 2.8 21 15 ns
6 0 3 3 85 3.1 9 24s
4 0 4 4 65 2.7 46 34 ns
6 0 0 6 50 2.6 3

Treated Females
0 8 2 2 28 1.3 15 14 ns
0 6 3 3 34 1.9 13 22s
0 4 4 4 40 2.7 32 34ns
0 6 6 0 56 2.9 1

0 0 6 6 46 2.9 66

*T = Treated; U = Untreated.
**Computed by methods of Fried (1971); X2 was computed for percentage observed vs. percentage
expected hatch.

the treated male cross and 3% in the treated female cross as compared with
74% for the checks.
Our favorable results for the heteroaromatic aziridines and the effec-
tiveness of organophosphorus aziridines against the tobacco budworm suggest
that we test related aziridines for even greater activity. We also need a method
to treat large numbers of insects with the effective chemosterilants. Topical
applications are not satisfactory. Perhaps the fumigation method of Chang et
al. (1973) could be developed for this purpose.


Bankwith, Jr., K. B., and J. B. Graves. 1970. Evaluation of selected
chemosterilants against the tobacco budworm. J. Econ. Ent.
Bowers, W. S. 1968. Juvenile hormone: Activity of natural and synthetic
synergists. Science (Wash. D.C.) 161:895-897.
Chang, S. C., A. B. Borkovec, C. W. Woods, and P. H. Terry. 1973.
Chemosterilization of male house flies by fumigation. J. Econ. Ent.

Vol. 57, No. 3, 1974

Wolfenbarger et al.: Sterilants for H. virescens

Flint, H. M., and E. L. Kressin. 1969. Transfer of sperm by irradiated Heliothis
virescens (Lepidoptera: Noctuidae) and relationship to fecundity. Can.
Ent. 101:500-507.
Flint, H. M., W. Klassen, E. Kressin, and J. Norland. 1968a. Chemosteriliza-
tion of the tobacco budworm: A survey of 21 compounds applied
topically. J. Econ. Ent. 61:938-941.
Flint, H. M., W. Klassen, E. Kressin, and J. Norland. 1968b. Chemosteriliza-
tion of the tobacco budworm: A survey of 16 compounds fed to adult
moths. J. Econ. Ent. 61:1726-1729.
Fried, M. 1971. Determination of sterile-insect competitiveness. J. Econ. Ent.
Guerra, A. A. 1970. Effects of biologically active substances in the diet and
development and reproduction of Heliothis spp. J. Econ. Ent.
Guerra, A. A., D. A. Wolfenbarger, and R. D. Garcia. 1973. Activity of
juvenile hormone analogues against the tobacco budworm. J. Econ.
Ent. 66:833-835.
Guerra, A. A., D. A. Wolfenbarger, D. E. Hendricks, R. D. Garcia, and J. R.
Raulston. 1972. Competitiveness and behavior of tobacco budworms
sterilized with reserpine and gamma irradiation. J. Econ. Ent.
Rudesill, J. T., R. F. Severson, and J. G. Pomonis. 1971. The synthesis of
N-arylaziridines. J. Org. Chem. 36:3071-3076.
Soto, P. E., and J. B. Graves. 1967. Chemosterilization of bollworms and
tobacco budworms. J. Econ. Ent. 60:550-553.
Wolfenbarger, D. A., and A. A. Guerra. 1971. Response of strains and sexes of
the tobacco budworm to gamma irradiation. J. Econ. Ent.
Wolfenbarger, D. A., and A. A. Guerra. 1972. Sexual competitiveness of
Co"o-irradiated tobacco budworms. J. Econ. Ent. 65:1596-1600.
Wolfenbarger, D. A., A. A. Guerra, and S. H. Robinson. 1973. Tobacco bud-
worms: Sterilization, sexual competitiveness, and sperm transfer of
moths reared from eggs irradiated with "6Co. J. Econ. Ent. 66:1067-1070.
Wolfenbarger, D. A., and W. L. Lowry. 1969. Toxicity of DDT and related
compounds to certain lepidopteran cotton insects. J. Econ. Ent.

The Florida Entomologist

CURCULIONIDAE)'-(Note.) The 5 major varieties of citrus rootstocks
grown in Florida, rough lemon, sour orange, 'Carizzo' citrange, 'Milam' rough
lemon, and 'Cleopatra' mandarin were tested for resistance to damage by
Diaprepes abbreviatus (L.). In August 1971, 10 trees of each variety, 1.9-2.5 cm
diam calipered 8 cm above soil level, and ca. 60.9-76.2 cm tall, were placed in
individual 208-liter steel drums with both ends removed and planted in a
double Latin square design. The drums were buried in the soil with ca. 3.8 cm
extending above ground level.
In September 1971 the 50 trees were each infested with 50 D. abbreviatus
larval 1st instars from the Apopka laboratory by dropping the larvae into a
hole ca. 4-5 cm deep in the soil adjacent to the tree trunk and filling the hole up
with soil. In April 1972, 5 trees of each rootstock were dug up and the soil in
each drum was washed through a series of screens to recover immature
weevils. Also, larval feeding injury to the roots of each tree was numerically
rated. Screens were fitted to the tops of the remaining drums to prevent the
escape of emerging adults, and each drum was checked daily for adult emer-
gence. In November 1972, the remaining trees were dug and evaluated as
No differences in resistance were observed. At the 7-month sampling, 14
larvae and 3 pupae were recovered. At 14 months, 1 larva was recovered. The
larvae were recovered at depths ranging from 30-79 cm; the pupae were at
depths of 2.5-20 cm. No adults emerged. Severe damage was done to the root
systems of the trees within 6 months, September to April. Evidently larval
activity continues during winter months. P. A. Norman, A. G. Selhime, and R.
A. Sutton2 U. S. Horticultural Research Laboratory, 2120 Camden Road,
Orlando, Fla. 32803.

'Received for publication 1 April 1974.
'The authors gratefully acknowledge the technical assistance of D. J. Lyon and W. McCloud of


Vol. 57, No. 3, 1974

The Florida Entomologist



Agricultural Research Service, USDA


The maximum tobacco budworm, Heliothis virescens (F.), population on
St. Croix occurs during October and November, while the low is during June,
July and August. Pigeon pea and Bastardia serve as the principle host plants
throughout the year. Population estimates show the species at a sufficiently
low level to make St. Croix an excellent location for suppression or eradication

St. Croix, a member of the U. S. Virgin Islands, is situated in the Caribbean
Sea 60 miles southeast of Puerto Rico. It has an area of 84 miles2 and offers
diverse ecological situations seldom found on a small land mass. Stanley et al.
(1971) gave detailed information concerning the physical features and en-
vironment. Since St. Croix is so isolated and thus potentially suitable for
evaluation of suppression and eradication techniques against the tobacco
budworm, Heliothis virescens (F.), a population survey was conducted there
from October 1970 to September 1971. In addition, considerable host plant
data have been collected, as well as a population estimate.

The survey system involved the periodic operation of sticky traps baited
with virgin female H. virescens as described by Snow and Copeland (1969).
These traps were constructed of quart plastic containers that had openings in
each end and were coated inside with a thin layer of StickemR. From 4 to 5
virgin females were held within the trap in smaller plastic cages and provided
with a small piece of sponge soaked in sugar water. A total of 28 sites was
selected for traps and all types of terrain were represented. The traps were
placed at 4-ft elevations on fences or stakes and were serviced twice weekly by
replacing the entire trap unit; thus, the bait females were usually in the field
for 3 or 4 nights only.

Lepidoptera: Noctuidae.
2Accepted for publication 13 Feb. 1974.
Mention of a commercial or proprietary product in this paper does not constitute an en-
dorsement of this product by the USDA.
Tobacco Insects Laboratory, St. Croix, U. S. Virgin Islands 00850.
Present address: Vegetable Germplasm Development Laboratory, Beltsville, Maryland 20705.
Tobacco Insects Laboratory, Oxford, North Carolina 27565.
Present address: University of Tennessee, Dep. of Engineering, Knoxville, Tennessee 37901.
Cotton Insects Research Laboratory, Brownsville, Texas 78520.

Vol. 57, No. 3, 1974


298 The Florida Entomologist Vol. 57, No. 3, 1974

In the release and recapture experiment, we used males reared on meridic
diet at Brownsville, Tex. and shipped as pupae to St. Croix. Calco oil red
N-1700R dye had been incorporated into the larval diet so the moths were
marked internally (Hendricks and Graham 1970). The newly emerged adults
were collected daily, cooled at 500F for 30 min, and placed in pre-chilled
insulated jars for irradiation (32 krad). After irradiation, the moths were
placed in 1-gal cages (50 males per cage) and held outdoors until late evening
at which time 50 males were released at each of 10 locations. All points on the
island were within a 2-mile radius of a release site. The release of 500 males per
night was made for 7 consecutive nights during which no traps were operated
(to allow the number of marked insects to become stabilized). At the end of the
week, 100 sticky traps, 15 grid traps, and 50 light traps were placed in opera-
tion. Daily releases of 500 males per day were then continued for 4 more days
until all 3 trap systems were operated and the capture recorded. Thereafter,
no releases were made, and the grid traps were operated daily to determine
how long the marked insects remained in the environment.
The sticky traps were of the type and operated in the same manner as in
the survey; the light traps were of the type described by Stanley et al. (1964);
and the grid traps were of the type described by Kishaba et al. (1970) with
slight modifications. The charged rods were spaced /2 in. apart in a cylindrical
pattern with 4,000 v AC applied across the elements. Ten virgin females were
used as bait. All 3 types of traps were placed randomly over the island, with
independent randomization of each type.
Suspect host plant species have been examined for tobacco budworm eggs
and larvae since 1967. After finding larvae on a plant periodic samples were
made to determine the infestation level and relative importance of the plant
species in the population dynamics of the insect.

Population Survey
The data for the survey have been combined into 2-week averages which
are reported in Table 1 as the average capture/trap-night. A shortage of virgin
females often prevented continuous operation so the actual number of nights
trapped within each 2-week period are given. The population appeared to be at
a maximum at the initiation of the trapping in October 1970. The lowest
population occurred during June and July 1971, probably because of an ex-
treme drought. All 28 traps caught males at some time, but more were cap-
tured in the arid eastern areas.
Host Plants
The host plant situation on St. Croix is complex, and includes several
species of plants and ovipositional patterns by the insects.
Pigeon pea, Cajan cajan (L.), appears to be the most important budworm
host. The plant is a bushy branched perennial growing up to 2 m high and is a
favored food by Virgin Islanders. Pods and blossoms can be found at all times
of the year and oviposition is usually on the pod. After eclosion from the eggs,
there is some feeding on the outside of the pod by small larvae but they soon
eat into the pod and consume the beans. Thirteen eggs and 1 larva/pod have
been the maximum infestation. The normal level has been about 0.05 larvae
and 0.20 eggs/pod. Tobacco grown for experimental reasons near pigeon pea
has shown lower egg and larval population levels than the adjacent peas.

Snow et al.: The Tobacco Budworm on St. Croix


Average capture/
Time period Total days trapped Trap/Day

Oct. 12-25, 1970 11 1.86
Oct. 27-Nov. 8 8 0.71
Nov. 9-22 0
Nov. 23-Dec. 6 3 1.02
Dec. 7-20 3 0.21
Dec. 21-Jan. 3, 1971 14 0.22
Jan. 4-17 7 0.58
Jan. 18-31 11 0.11
Feb. 1-14 13 0.24
Feb. 15-28 14 0.63
Mar. 1-14 14 0.16
Mar. 15-28 14 0.11
Mar. 29-Apr. 11 0
Apr. 12-25 14 0.39
Apr. 26-May 9 14 0.07
May 10-23 10 0.27
May 24-June 6 14 0.28
June 7-20 8 0.09
June 21-July 4 12 0.01
July 5-18 0
July 19-Aug. 1 7 0.11
Aug. 2-15 4 0.12
Aug. 16-29 4 0.43
Aug. 30-Sept. 12* 4 0.51

*Based on captures in 100 sticky traps.

The malvaceous plant, Bastardia viscosa (Kth.), appears to be the second
most important budworm host on St. Croix and infestations often exceed
those found on pigeon pea. This plant is fairly pubescent and very sticky to the
touch. The plant occurs throughout St. Croix but mostly in the flat areas
receiving 25-40 in. of rain. The plant is regularly attacked, with most oviposi-
tion and feeding on the upper leaves and fruit.
Turkey berry, Solanum torrum Sw. Prodr., a rather high and stout
branched plant with fruit about 10 to 14 cm diam, is fairly abundant in the
northwestern hills. The maximum attack level has been 8 eggs and 0.5 lar-
vae/flower head. However, population levels are usually below this level and
based on abundance and the sporadic nature of infestations, the plant proba-
bly is of importance only at certain times of the year.
Beggar's lice, Desmodium spp., is a perennial herb sparsely found in low to
high elevations throughout the 30-60 in. rainfall area of St. Croix. Oc-
casionally the plant has been found heavily infested with eggs and larvae.
However, in terms of budworm survival on St. Croix the plant is probably of
little importance.


The Florida Entomologist

Other wild plants that are occasionally attacked on St. Croix are Pop
Bush, Passiflora foetida L., and Spider-Flower, Cleome spinosa Jacq. Enum.
Other wild host species probably exist that have not been found. Populations
of the tobacco budworm have been found on the pods of okra when it is
available. Generally pods are harvested while larvae are small and the acreage
is extremely limited. Therefore, the plant probably contributes little to the
island budworm population.
Population Estimate
The results of the release and recapture experiment are shown in Tables 2
and 3. Table 2 reports the capture in the 3 trap systems from 3 to 7 Sept., a
period during which daily releases were being made. During the 4 nights the 15
grid traps caught over twice as many native and released males as the sticky
traps. Only 1 native male was positively identified from a light trap. The large
number of other species caught in light traps made identification difficult,
though the inefficiency of these traps compared with grid and sticky traps was


Capture Ratio
Type trap* Marked Native Marked:Native

50 light traps 0 1 -
15 grid traps 91 473 1:5.20
100 sticky traps 27 202 1:7.48
Totals 118 676 1:5.73

*Sticky traps baited with 5 virgin females and grid traps with 10 virgin females.

As shown in Table 2, a total of 118 released and 676 native males were
captured, a 1:5.73 ratio. Thus it would have required the release of 2865
males/day (instead of 500) to achieve a 1:1 ratio of natural to marked males.
To achieve ratios necessary to attempt suppression or eradication at this
population level (by the sterile male technique) 28,650 males per day would
have been required to achieve a 10:1 ratio and 57,300 males per day to achieve
a 20:1 ratio. However, if the releases had begun in June or July when sticky
traps were catching only 0.10 males per night (they were catching 0.51/night
at the time of the population estimates), presumably only 1/5 as many
released males would be needed. These estimates are valid for Brownsville-
reared insects treated and handled in the procedures used in our experimen-
tation. In this type of estimate, mortality factors, vigor of the released insects,
etc., are automatically incorporated into the population estimate.
The 676 native insects caught from 3 to 7 Sept. were taken from all areas of
St. Croix ranging from the drier eastern tip (less than 25 in. rainfall/year) to

Vol. 57, No. 3, 1974

Snow et al.: The Tobacco Budworm on St. Croix

the rain forests in the northwestern hills (average 60 in. rainfall/year). Ap-
parently, the species is able to infest the entire island.
The captures in the 15 grid traps are reported by dates in Table 3. The last
release of marked males was made 7 Sept., but marked males were recovered
for the next 8 days. This daily operation of the grid traps was intended to
demonstrate the daily loss of released insects in the environment. Virgin
females are not equally attractive each night so this evaluation can not
be considered valid. Nevertheless, the fact that these released males sur-
vived for 8 nights is important. All indications show St. Croix as an excel-
lent location for suppression or eradication studies for the tobacco bud-


Date operated Age of bait (days) Native Marked

Sept. 3-7 1-4 473 91
Sept. 8 1 103 29
Sept. 9 2 230 40
Sept. 10 3 139 14
Sept. 11 1 168 14
Sept. 12 2 180 8
Sept. 13 3 122 9
Sept. 14 1 158 6
Sept. 15 2 202 2
Sept. 16 3 178 0


Hendricks, D. E., and H. M. Graham. 1970. Oil-soluble dye in larval diet for
tagging moths, eggs, and spermatophores of tobacco budworms. J.
Econ. Ent. 63:1019-20.
Kishaba, A. N., W. W. Wolf, H. H. Toba, A. F. Howland, and T. Gibson. 1970.
Light and synthetic pheromone as attractants for male cabbage
loopers. J. Econ. Ent. 63:1417-20.
Snow, J. W., and W. W. Copeland. 1969. Fall armyworm: Use of virgin female
traps to detect males and to determine seasonal distribution. USDA
Prod. Res. Rep. 110, 9 p.
Stanley, J. M., F. R. Lawson, and C. R. Gentry. 1964. Area control of tobacco
insects with blacklight radiation. Trans. Amer. Soc. Agr. Eng. 7:125-7.
Stanley, J. M., A. H. Baumhover, W. W. Cantelo, J. S. Smith, Jr., M. B. Peace,
and C. A. Asencio. 1971. A population suppression experiment for
tobacco hornworms and other nocturnal insects using blacklight traps
on an isolated island, preliminary studies. USDA ARS 42-193. 8 p.

The Florida Entomologist

Abstract.) Fifty of each sex of 2 species of terrestrial isopods, Armadillidium
vulgare and Porcellio laevis, were each offered 16 live pupae, 4 each of
Drosophila hydei, D. immigrants, D. pseudoobscura, and D. melanogaster, for
24 hr. Both sexes of both predator species fed on all prey species, each predator
species taking numbers of prey species in the reverse order of prey size, namely
D. melanogaster > D. pseudoobscura > D. immigrants > D. hydei. The
results, however, do not indicate preference by predators. In terms of total
weights consumed, Porcellio males and females ate 8.91 and 8.04, and Ar-
madillidium males and females 11.69 and 11.86 mg/isopod/day, respectively.
The difference between sexes is insignificant, but between predators it is
highly significant (P < 0.001). Rates of consumption of Drosophila pupae and
of Hippelates pupae decreased by about two-thirds or more over a week in the
absence of other food.
Laboratory experiments using known numbers of :1P-labelled Drosophila
pupae fed to Armadillidium proved the feasibility of using this technique for
field studies. Such work subsequently showed that A. vulgare ate D.
melanogaster pupae in natural conditions in a citrus grove where alternative
food was abundant. If predation by isopods in nature proves to be wide-spread,
their significance in several ecosystems would deserve reconsideration.
(Ecology, 1974, 55(2):428-433; E. B. Edney, Univ. Calif., Riverside, 92502, W.
Allen, Univ. Calif., Los Angeles, 90024, and J. McFarlane, Univ. Calif., River-
side, 92502).

Abstract.) Mark-recapture techniques were used to study the immigration
and emigration of adults of the Queensland fruit fly in an isolated orchard at
Wilton, N.S.W. During the 1968 season, several thousand mature flies
migrated into the orchard during late February and early March; the females
in this influx were responsible for laying the majority of eggs which con-
tributed to the next generation. After the fruit disappeared these immigrants
left the orchard, but immature flies continued to enter the orchard in smaller
numbers until late in the season. In the early part of the season approximately
75% of adults that emerged in the orchard left during their first week. Later in
the season, when lack of rain made conditions less favorable, nearly all the
flies left the orchard in the first week after emergence.
During the 1969 season, when no fruit was present because of a spring
drought, fewer mature flies migrated into the orchard in late February and
March, even though traps located over a wide area around the orchard in-
dicated that at least as many flies were present in the general area as in 1968.
Studies on emigration rates at the orchard were carried out using flies bred
from infested fruit collected in neighboring towns. As in the comparable part
of the 1968 season, approximately 75% of the flies left the orchard in the first
week after emergence. It is postulated that dispersive movements in D. tryoni
can be divided into 3 main categories: a post-teneral dispersive stage, host
seeking, and responses to adversity. (Aust. J. Zool., 1973, 21(4):541-65; B. S.
Fletcher, Univ. Sydney, N.S.W. 2006).

Vol. 57, No. 3, 1974

The Florida Entomologist



Multiple regression models of 11 species of adult insects representing the
orders Anoplura, Coleoptera, Diptera, and Lepidoptera indicated that total
body concentrations of several major and trace elements could be used as a
bioindicator or biopredictor of radiation-induced insect sterility on a species
level. There was a particularly high correlation between total body major and
trace element content and irradiation-induced insect sterility in a group of
stored-product insects that had similar feeding preferences (i.e., laboratory
diets of grain and/or flour).
A statistical model based on insect total body Cu and K/Mg ratio was
highly effective in estimating or predicting the radiosterilization doses for
several species of stored product insects feeding primarily on grain and/or

Total body concentrations of several major and trace elements cationss)
have been shown to be correlated to the acute (LD50/24 hr) radiosensitivity of
stored product beetles, cockroaches, and blood feeding insects (Levy et al.
1973). Simple regression analyses indicated that Cu or Cu/Fe ratio, K, and Mg
were the most effective biological predictors or indicators of acute radiation
mortality for the 3 groups respectively. Additional research utilizing multiple
regressison analyses indicated the interelement dependence of Na with K (i.e.
Na + K) and Mg (i.e. Na + Mg) in formulating predictor models for estimating
the acute radiation mortality of cockroaches and blood feeding insects re-
spectively (Levy et al. in press).
The dietary and metabolic importance of major and trace elements have
been briefly reviewed by Levy and Cromroy (1973). Englemann (1970)
presented an excellent review showing the importance of dietary concentra-
tions of Zn, Fe, K, Mg, Na, and Cu in the reproductive physiology of adult and
immature insects.
The aim of this research was to determine whether total body concentra-
tions of several major and trace elements can be used as an effective biological
indicator or predictor of radiation-induced sterility on a species level.
Predictor equations could, therefore, be used to expedite the procedures
involved in determining a species-specific radiosterilization dose for insect
control, e.g. in a sterile-male release program. Such information could also be
used to evaluate insect and other biological imbalances produced in a post-
nuclear attack environment.
The total body concentration of Cu, Fe, K, and Mg in 11 species of
adult insects representing the orders Anoplura, Coleoptera, Diptera, and

Florida Agricultural Experiment Station Journal Series No. 5095.
2Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611.
: Department of Statistics, University of Florida, Gainesville, Florida 32611.

Vol. 57, No. 3, 1974

The Florida Entomologist

Lepidoptera were obtained from the atomic absorption spectrophotometric
data of Levy and Cromroy (1973). Sterilization doses for adult insects were
obtained from the radiation literature (International Atomic Energy Agency
Bibliographical Series Nos. 9(1963), 15(1965), 24(1967); Stern (1969) from
samples of mixed sexes or obtained separately for males and females and
averaged to give a general sterilization dose for an insect species population.
The sterilization dose for a species was defined as that dose in rads (radiation
absorbed dose) of x or gamma radiation required to induce 99.9-100.0% sterility
in the irradiated insects. Sterilization doses obtained in units of R (Roentgen)
were converted to rads by multiplying by a conversion factor of 0.93.
Statistical analyses consisted of fitting multiple regression models using
the method of least squares (Draper and Smith 1966). These analyses were
based on the straight line equation, y=b,+b,(X,)+b,(X,), where y=es-
timated or predicted radiosterilization doses in rads, b,,= constant,
b,...b, = estimate or measure of the strength of X,...X, = total body concentra-
tion of a specific element or ratio of elements in parts per million (ppm).
To test the strength of a predictor equation, the coefficient of multiple
determination (R2) and F-statistic (F) were calculated for each model. F-test
significance at the 0.05 level was indicated for each model. The standard error
(SE) for each predicted or estimated radiosterilization dose was also deter-

Results from multiple regression analyses indicated that a model
representing the ratio of 2 trace (Cu/Fe) and 2 major (K/Mg) elements was
the most effective indicator or predictor of radiation-induced sterility for a
wide range of insect species (Table 1).
Relationships between trace (Cu/Fe) and major (Na & Mg, Na & K)
elements have been shown to be useful in estimating or predicting the
radiosensitivity of stored product beetles, cockroaches, and blood feeding
insects (Levy et al. 1974). A biological relationship has been shown between
Na and Mg, hence, it could be conjectured that a relationship exists between
K and Mg in insects (Chapman 1969).
Although the predictor equation (Table 1) was significant at the 0.05 level,
it only accounted for 75% (i.e. R = 0.7478) of the observed radiosterilization
doses. This was assumed to be due primarily to variations in diet, since Levy et
al. (1973) have shown a relationship between radiosensitivity and insect die-
tary (i.e. the total body concentration of major and trace elements ac-
cumulated from an insect diet). Additional variations between the observed
and predicted radiosterilization doses were presumed to be due to phylogene-
tic relationships, body weight and size, and physical activity (Menhinick and
Crossley 1969). Therefore, the insect species were subdivided and an analysis
made of a group that had similar feeding preferences, i.e., stored-product
insects fed grain and/or flour.
Multiple regression analyses of 7 species of stored product insects
representing the orders Coleoptera and Lepidoptera suggested an
interelement dependence of major and trace elements in formulating effective
models for predicting insect radiosterilization (Table 2). The data indicated
that total body concentrations of several biologically active trace (Cu) and
major (K/Mg) elements cationss) could be used as an excellent biological
indicator of predictor of radiation-induced sterility of stored product insects


Vol. 57, No. 3, 1974

Levy et al.: Predictors of Insect Sterility



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0 c^


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Levy et al.: Predictors of Insect Sterility 307

on a species level when insects were subdivided into a general feeding group
based on laboratory diets consisting mainly of grain and/or flour. Levy et al.
(1973) have shown the importance of Cu in estimating or predicting the acute
radiosensitivity of stored product beetles.
The removal of Fe from the stored product insect model seemed to
eliminate an erroneous feeding variable accumulated from an insect diet not
having mineral levels comparable with a diet of grain and/or flour. This
subsequently increased the prediction capability by significantly (i.e.
R = 0.9666) decreasing the variation between the observed and predicted
radiosterilization doses.
These data suggest the physiological importance and interaction of certain
trace and major elements cationss) in the mechanisms) involving the
radiosensitivity of insects. In addition, the stored product insect grouping was
presumed to have reduced phylogenetic variations, as well as variations in
body size, weight, and physical activity which have been shown to be related to
insect radiosensitivity.

This research was partly supported by NIH Training Grant No.
1T01A100383-01 from the National Institute of Allergy and Infectious


Chapman, R. F. 1969. The Insects: Structure and Function. American
Elsevier Publishing Co., Inc. 819p.
Draper, N. R., and H. Smith. 1966. Applied regression analysis. John Wiley &
Sons, Inc., N.Y. 407p.
Englemann, F. 1970. The physiology of insect reproduction, Vol. 44. Pergamon
Press, N.Y. p. 117-118.
Levy, R., and H. L. Cromroy. 1973. Concentration of some major and trace
elements in forty-one species of adult and immature insects determined
by atomic absorption spectroscopy. Ann. Ent. Soc. Amer. 66:523-26.
Levy, R., H. L. Cromroy, and J. A. Cornell. 1973. Major and trace elements as
bioindicators of acute insect radiosensitivity. Radiat. Res. 56:130-39.
Levy, R., H. L. Cromroy, and J. A. Cornell. 1974. Multi-elemental models for
estimating the acute radiosensitivity of cockroaches and blood feeding
insects. Fla. Ent. 57:43-46.
Radioisotopes and Ionizing Radiations in Entomology, Vol. I, (1950-1960),
Bibliographical Series No; 9, p. 371. International Atomic Energy
Agency, Vienna, 1963.
Radioisotopes and Ionizing Radiations in Entomology, Vol. II, (1961-1963),
Bibliographical Series No. 15, p. 435-37. International Atomic Energy
Agency, Vienna, 1965.
Radioisotopes and Ionizing Radiations in Entomology, Vol. III, (1964-1965),
Bibliographical Series No. 24, p. 326-33. International Atomic Energy
Agency, Vienna, 1967.
Stern, V. M. 1969. Insect pests of major food crop, their reinvasion potential
and the effects of radiation on arthropods. OCD Work Unit No. 3145B,
University of California.

The Florida Entomologist

TIDAE)-(Note.) During the spring of 1973, while making routine insect
inspections and collections in various grasses, the senior author observed
numerous deadhearts in stems of vaseygrass, Paspalum urvillei Steud. Stem
examination showed these to be infested internally with dipterous larvae and
puparia. The stems were attacked prior to emergence of the seedhead, i.e.,
during the boot stage of development. The newly hatched maggots severed the
pre-emergent seedhead and fed on the aborted inflorescence, resulting in a
rather moist necrotic situation so often typical of maggot activity. Outwardly
the sign of attack is hardly distinguishable from that of the curculionids
Centrinaspis picumnus Herbst and Centrinaspis sp. attacking various digit
grasses, Digitaria spp., and other grasses.
Several larvae and pupae were collected for rearing to the adult stage. The
adult insects were brown, rather elongate, picture-winged (otitid) flies. The
junior author determined these to be the species Zacompsia fulva Coquillet.
Inspection did not indicate infestation in any other grasses. Since P. ur-
villei usually is considered a weed grass in Florida, no economic damage was
attributed to this fly. However, since 20 to 40% of stems were infested and since
related grasses, particularly bahiagrass, P. notatum Flugge, are grown for
pasture and turf in the Everglades and adjacent areas, the fly will bear
watching in situations involving seed production. No previous Z. fulva host
report was found. C. H. Curran (1934. The Families and Genera of North
American Diptera; Ballou Press N. Y.) mentioned Zacompsia but only in his
key to the genera. Alan Stone et al. (1965. A Catalog of the Diptera of America
North of Mexico; USDA Agr. Handbook No. 276. Washington, D. C.) recorded
the range as: TX and LA to SC south to FL. K. Valley et al. (1969. Ann. Ent.
Soc. of America 62:227-34) reported an otitid of similar habits, Eumetopiella
rufipes (Macquart), attacking inflorescences of barnyard-grass, Echinochloa
crusgalli L. (Beauv.), in NY and OH. This was the only previous instance
reviewed by Robert Lavigne (1974. Bull. Ent. Soc. of Amer. 20(1):11-23) of an
otitid attacking inflorescences of living grasses, but grain crops were excluded
from the review. William G. Genung, Agr. Res. and Educ. Center, Belle Glade,
FL. 33430 and Howard V. Weems, Jr., FL Dep. of Agr. and Consumer Services,
Div. of Plant Indus., Bur. of Ent., Gainesville, FL 32602.

Vol. 57, No. 3, 1974

The Florida Entomologist


Stored-Product Insects Research and Development Laboratory,
Agr. Res. Serv., USDA, Savannah, Georgia 31403

The flat grain beetle, Cryptolestes pusillus (Schonherr), and C. turcicus
(Grouvelle) were reared at a high temperature (29 1C) on 20 diets prepared
from a single natural product or mixture of natural products. Wheat was the
most favorable natural diet and cracked food was the most favorable form of
diet. Neither C. pusillus nor C. turcicus multiplied on brown rice meal, runner
peanuts, whole soybeans, cracked black-eyed peas or Purina Chow''R with
brewer's yeast; brewer's yeast was not significantly advantageous in mixed
diets. Cryptolestes pusillus multiplied more on all diets except cracked
soybeans and wheat meal than did C. turcicus.

The flat grain beetle, Cryptolestes pusillus (Schbnherr), and C. turcicus
(Grouvelle) are widely distributed, abundant, and economically important
species (USDA 1965) that infest a variety of stored commodities (Davies 1949,
Howe and Lefkovitch 1957). Information on the extent of damage caused by
these insects is generally available (Cotton 1963), but no data are available
concerning the relative buildup of the populations on different commodities.
Because I was interested in the population increase of the 2 species in the
South, I determined the increases in their populations at a high temperature
on 20 diets prepared from a natural product or a mixture of natural products.

Cryptolestes turcicus obtained from Canada were initially reared as a stock
culture at the Savannah laboratory on a mixture of white flour, white corn-
meal, and brewer's yeast (10:10:1.5 by vol). Then 4 months before the
experiment, the colony was placed on a mixture of cracked, soft red winter
wheat, rolled oats, and brewer's yeast (14:14:1.5 by vol), the standard diet used
at this laboratory to rear the stock cultures of C. pusillus. The test was begun
4 months later when both species had exhibited similarly excellent population
growth without excessive mortality on this diet. The procedure was similar to
that described for an earlier test (LeCato and McCray 1973): 5 pairs of 1-wk-
old adults of each species were placed into 1-pint (ca. 0.47-liter) jars on 200 ml
of each of 20 test diets. The jars were capped with a No. 1 filter paper disc
secured with a screw-type ring.

Coleoptera: Cucujidae.
Mention of a commercial or proprietary product in this paper does not constitute an en-
dorsement of this product by the U. S. Department of Agriculture.

Vol. 57, No. 3, 1974


The Florida Entomologist

The cracked corn, wheat, brown rice, soybeans, and black-eyed peas used
in the test diets were produced in a Viking'R hammer mill equipped with a
screen containing openings 3/8 in. (9.5 mm) in diam. To produce meal that
would pass through a U. S. Standard no. 30 sieve, a screen containing openings
1/64 in. (0.4 mm) in diam was used. Whole shelled runner peanuts were placed
in a commercial Waring(R) Blender (ca. 12,500 rpm) for 10 sec to produce
cracked peanuts and for 1 min to produce peanut meal.
Moisture content was determined with a Motomco Model 919'"' moisture
meter. Moisture content of corn, wheat, rice, and black-eyed peas was ad-
justed to ca. 12.5%. Moisture contents of soybeans and of peanuts were ad-
justed to ca. 12% and ca. 7%, respectively. All diets were held for 1-2 wk prior to
the experiment at the experimental conditions of 29+ 1 C, 65 +10% RH, and
12-hr light-dark cycles.
After 10 wk, the adult insects present in each jar were counted, and the
number of seeding adults (10) was subtracted to determine the number of
adult progeny. Each combination of diet and species of insect was replicated 5
times. Differences in multiplication were assessed by using analysis of
variance and Duncan's multiple range test.

In general, wheat was the most favorable diet, and cracked food, which
provided better harborage and oviposition sites, was the most favorable form
of diet (Table 1). Cryptolestes pusillus multiplied on more diets and produced
more progeny on each diet (except cracked soybeans and wheat meal) than C.
turcicus. Neither species multiplied on rice meal, any form of runner peanuts,
whole soybeans, cracked black-eyed peas, or Purina Chowf"' with brewer's
yeast. Rice and soybeans in any form were generally unfavorable to both
species, and neither could penetrate the seed coat of whole soybeans. Neither
species increased on peanuts though Cryptolestes spp. do infest peanuts (Howe
and Lefkovitch 1957); oil from peanuts probably limits any population
increase (Thomas and Shepard 1940). The seeding adults of both species
generally died on diets that were not advantageous to multiplication.
Both species produced relatively few progeny on whole grain, probably
because of the small size of the adult Cryptolestes (ca. 1/16 inch or ca. 1.6 mm)
and the relative weakness of the mouthparts (Cotton 1963). For example,
Cryptolestes pusillus was reported unable to attack undamaged grain (Payne
1946, Davies 1949) and multiplied slowly on diets of whole corn, wheat, and rice
(rice = 2 +1.7); apparently the germ of slightly damaged kernels served as the
source of food. Cryptolestes turcicus multiplied even more slowly on corn and
wheat and not at all on rice. Diets of mixtures of natural products plus
brewer's yeast did not significantly increase the populations of either species.
The temperature at which the test was conducted may have given an
advantage to C. pusillus. Currie (1967) reported that C. pusillus developed
faster than C. turcicus when the temperature was above 250C but that the
growth of populations of C. pusillus is limited by low temperature and low
relative humidity. C. pusillus is most abundant in the wet tropical and warm
temperate regions where it increases rapidly. Cryptolestes turcicus is more cold
hardy than C. pusillus (Solomon and Adamson 1955) and is found in the moist
temperate regions of the world (Howe and Lefkovitch 1957). The findings
reported here and the literature (Eden 1967) indicated that C. pusillus is the
more economically important species in the southern United States.

Vol. 57, No. 3, 1974

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