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(ISSN 0015-4040)
FLORIDA ENTOMOLOGIST
(An International Journal for the Americas)
Volume 66, No. 2 June, 1983
TABLE OF CONTENTS
The 66th Annual Meeting of the Florida Entomological Society
RICHMAN, D. B., W. H. WHITCOMB, AND W. F. BUREN-Predation on
Neonate Larvae of Diaprepes abbreviatus (Coleoptera: Cur-
culionidae) in Florida and Puerto Rico Citrus Groves ............ 215
GERSABECK, E. F., AND R. W. MERRITT-Vertical and Temporal Aspects
of Alsynite@ Panel Sampling for Adult Stomoxys calcitrans
(L.) (Diptera: M uscidae) ..................................................-.......... 222
LINLEY, J. R.-Autogeny in the Ceratopogonidae: Literature and
N otes ......----. ------....... ..................................................................... 228
MORGANTE, J. S., A. MALAVASI AND R. J. PROKOPY-Mating Behavior of
Wild Anastrepha fraterculus (Diptera: Tephritidae) on a
Caged Host Tree .....--------... ------.................. ...................... ................. 234
MOCKFORD, E. L.-Systematics of Asiopsocidae (Psocoptera) Including
Pronotiopsocus amazonicus N. Gen. N. Sp. ..........................-......... 241
KRASNOFF, S. B., K. W. VICK, AND R. W. MANKIN-Female Calling
Behavior in Ephestia elutella and E. figulilella (Lepidoptera:
Pyralidae) ................. ...................................................-------............. 249
PECK, S. B.-New Cavernicolous Proptomaphaginus from Hispaniola
and Mexico (Coleoptera: Leiodidae: Cholevinae) ........................ 254
ASHLEY, T. R.-Growth Pattern Alterations in Parasitized Fall Army-
worm Larvae ................--- .......--.......................-........................ ........... 260
ASHLEY, T. R., et al.-Parasitization of Fall Armyworm Larvae on
Volunteer Corn, Bermudagrass, and Paragrass ............................ 267
BIEMAN, D. N., AND J. A. WITTER-Mating Behavior of Malacosoma
disstria at Two Levels of Mate Competition ..............................-..... 272
Scientific Note
HARPER, J. D.-Late Season Soybean Looper (Lepidoptera:
Noctuidae) Population Reduction by Western Palm
W arblers ........--- .........--................................ ............--.............- 280
Book Notices ...----. ------.............. -.........--..----...-................... ...................... 282
Smithsonian Foreign Currency Program .............. .......--- ...- ................. 283
Sustaining Members .........-.. ............----------------............-..... 284
FLORIDA ENTOMOLOGICAL SOCIETY
OFFICERS FOR 1982-83
President ....----- -----------........................................................... A. C. (Abe) W white
President-Elect .---...............-----.-.....................-- ................... C. W McCoy
Vice-President ----........----------..- -.....-.... -........- ......-. M. L. Wright, Jr.
Secretary ...---............-.............. ---...................................___ D. F. Williams
Treasurer .........---.-----.........--- .............-----................-- .___ D. P. Wojcik
J. R. Cassani
J. L. Knapp
D. C. Herzog
Other Members of the Executive Committee ... K. Lee
C. A. Morris
W. L. Peters
C. A. Musgrave Sutherland
PUBLICATIONS COMMITTEE
Editor ........................................ ................ ..- C. A. Musgrave Sutherland
Associate Editors ..----........ -----------------........... ....... D. C. Herzog
F. W. Howard
M. D. Hubbard
J. R. McLaughlin
A. R. Soponis
H. V. Weems, Jr.
Business M manager ............................................................................... D. P. W ojcik
FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September,
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Business matters for other Society officers can be sent to that individual at
the University Station address above.
This issue mailed June 27, 1983
THE 66th ANNUAL MEETING OF THE
FLORIDA ENTOMOLOGICAL SOCIETY
THIRD ANNOUNCEMENT
The Florida Entomological Society will hold its 66th Annual Meeting on
9-12 August 1983 at the Sheraton Sand Key Hotel, Clearwater Beach, Flor-
ida. The location is 1160 Gulf Boulevard, Clearwater Beach, Florida 33515;
telephone-1-(813)-595-1611. Room rates will be $45.00 either single or
double; student rates will be a flat $15.00 per person, with a minimum of
3 students per room.
Questions concerning the local arrangements should be directed to:
DR. ROBERT W. METZ, Chairman
Local Arrangements Committee
Florida Entomological Society
1808 North 57th Street
Tampa, Florida 33619
Phone: 1-(813)-626-3184
The deadline for submitting papers for the 1983 meeting was 1 June 1983.
Eight minutes will be allotted for presentation of oral papers, with 2
minutes for discussion. In addition, there will be a separate session for
members who may elect to present a Project (or Poster) Exhibit.
The three oral student papers judged to be the best on content and
delivery will be awarded monetary prizes during the meeting. Student
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Richman et al: Predation of Diaprepes Larvae
PREDATION ON NEONATE LARVAE OF DIAPREPES
ABBREVIATUS (COLEOPTERA: CURCULIONIDAE)
IN FLORIDA AND PUERTO RICO CITRUS GROVES
DAVID B. RICHMAN, W. H. WHITCOMB, AND W. F. BUREN
Department of Entomology and Nematology
University of Florida
Gainesville, FL 32611 USA
ABSTRACT
Predation by ants on neonate (1st instar) larvae of Diaprepes abbrevi-
atus (L.) on the soil surface in citrus groves was less in the afternoon
(1200-1530 EST) in Florida (average 9.6% in 2 groves) compared with
previously reported evening through morning periods (average 47% for 1
grove). Predation averaged 6.0% at Isabela, Puerto Rico from morning
through afternoon and 1.5% at Adjuntas, Puerto Rico in the afternoon
(1200-1600 PM AST). Researchers at the U.S. Dept. of Agriculture ARS
Lab at Plymouth, FL have shown that most larvae drop from the trees and
burrow into the ground from 1200-2000. It is thus concluded that in contrast
to the findings of earlier researchers, surface predation by ants is probably
much less important than other mortality factors, such as egg predators,
subterranean ants and nematodes, and climatic conditions.
RESUME
La rapacidad de las hormigas sobre las larvas neonatas (10 instar) de
Diaprepes abbreviatus (L.) en la superficie del suelo en los naranjales fue
menos por la tarde (1200-1530 EST) en Florida (promedio de 9.6% en 2
naranjales) en comparasi6n con los periods que se extendian de la tarde
hasta la mafiana, los cuales fueron reportados previamente (promedio de
47% para 1 naranjal). El promedio de la rapacidad desde la manana hasta la
tarde (1200-1600 PM AST) fue 1.5%. Los investigadores del laboratorio
del U.S. Department of Agriculture en Plymouth, FL han mostrado que la
mayoria de las larvas caen de los arboles y entran en el suelo desde las 1200
hasta 2000 h. Se concluyen que, en contrast a lo que se han informado los
investigadores anteriores, la rapacidad por las hormigas de la superficie
probablemente es menos important que otros factors de la mortalidad, tales
como los predadores de los huevos, las hormigas subterraneas, los nemato-
dos, y las condiciones climaticas.
The sugarcane rootstalk borer, Diaprepes abbreviatus (L.), poses a
serious threat to citrus, foliage, and possibly sugarcane industries in
Florida. The weevil was apparently introduced from Puerto Rico to the
area around Apopka in 1964 (Woodruff 1964). Its range has since expanded
within Florida (Woodruff 1968).
D. abbreviatus adults feed on foliage of citrus, mango, and many other
plants. After mating, the females deposit their eggs between host plant
leaves glued together with an adhesive produced by the female. The eggs
hatch almost exactly in 7 days and the larvae drop to the surface of the
ground and burrow into the roots of the host plant (Woodruff 1968, Whit-
comb et al. 1982).
Studies of the predation on D. abbreviatus by arthropods have been
215
Florida Entomologist 66 (2)
made for the eggs (Richman et al. unpublished data) and larvae (Whitcomb
et al. 1982, Tryon and Whitcomb unpublished data). Immature stages seem
to be most vulnerable to attack, although adults have been occasionally
preyed on in Puerto Rico by lizards (Wolcott 1924) and birds (Wetmore
1916, Whitcomb et al., personal observations), and in Florida by spiders
(Gowan and Whitcomb, personal observations) and snakes (Gowan, personal
observations). The predation data of Whitcomb et al. (1982) were taken
mostly in the late afternoon to early morning because the exact time the
weevil larvae left the egg masses was unknown. They report very high rates
for larvae artificially placed in orchards at night and in the early morning.
Recent work by Jones and Schroeder (1983) indicates that the larvae
actually drop to the soil surface between 1200 and 2400 EST. The number
of larvae dropping decreased considerably by 2000 EST and ended completely
by midnight. Whitcomb et al. (1982) found little predation at 1100 and
1400 EST at Forest City, FL. They found high predation at the same
site at 1700 and 2300 EST (40%+) and very high predation (62%) at
0600 EST. Jones and Schroeder (1983) have shown in laboratory ex-
periments that most larvae burrow into the soil within 3 h of dropping. Thus
it appears that neonate larvae may be escaping many of the natural surface
predators. To test this hypothesis further, data on the importance of preda-
tors during the afternoon was collected in Florida during the summer of
1981 and in Puerto Rico in May 1982.
MATERIALS AND METHODS
Two citrus groves within the quarantine area for D. abbreviatus, in
Florida were chosen for comparative observations. They each had trees with
uniform canopies that had not been heavily frost damaged and had a
diverse arthropod fauna. These groves were at Forest City, Seminole Co., and
Lockhart, Orange Co. Weevil larvae were obtained from the U.S. Depart-
ment of Agriculture ARS Laboratory at Plymouth. Initially 50 larvae were
placed in plastic petri dishes (Whitcomb et al. 1982) and starting at 1200-
1230 EST 2 dishes were placed on the ground beneath each of 3 arbitrarily
selected trees within the grove. An observer at each tree watched the larvae
constantly for a 20-minute period. Dishes with fresh larvae were then
placed under new trees after each 20-minute period. This was repeated so that
a total of 1 h of observations per observer was completed for each series. A
second set of observations was conducted starting at 1400-1430 and continued
to 1500-1530. A total of 1800 larvae were observed per day under this system.
This method was used during July 1981.
After late July there were 2 observers. Also, since plain petri dishes did
not simulate natural surfaces enough to keep the larvae from escaping
completely during hot afternoons, petri dishes filled with a mixture of
plaster of Paris and sterile soil were used. In this case 6, 1-h observation
periods were conducted in each grove for a total of 24 h of observation on
7200 larvae. No attempt was made to keep larvae from leaving the dishes.
However, they were more likely to remain within the dish than when un-
altered dishes were used. Predation was recorded by direct visual observa-
tion of each event. Samples of each species of predator observed were taken
to the laboratory for identification. Direct visual observation was used in
this study because early attempts to check observation after 20 minutes
216
June, 1983
Richman et al: Predation of Diaprepes Larvae
gave less accurate predation rates and allowed larvae to escape at a much
higher rate (50% or more than plain dishes were used) during the hot
afternoon periods than was observed by Whitcomb et al. (1982) during the
evening and morning. Observations utilizing this method were made from 31
August to 2 October 1981.
Weevils were present at Forest City from June through October and
field-caught females were mating and laying eggs at the U.S. Department
of Agriculture ARS Lab at Plymouth, during this time. Temperature, rela-
tive humidity, cloud cover and general conditions were recorded at the
beginning and end of each hour period. The observations were made from
1200-1330 and from 1400-1530 EST in Florida. The Florida groves were
sprayed with ethion and oil during the summer and fall and dicofol for mites
at various times. The Forest City grove also had been treated with heptachlor
as recently as 1975 and with benomyl, wettable sulphur and malation
prior to 1980. Both Florida groves had a high population of ants and spiders
on the surface of the soil. The weather at Forest City and Lockhart was
quite variable during the afternoon, but the temperature was often above
300C and the relative humidity was 60% or higher.
Observations in Puerto Rico were based on the same technique, except
that 3 observers with 1 dish each were utilized and observations were
conducted in the morning, at night and in the afternoon, for comparison
with earlier work in Florida. A total of 7.5 h of observations (3 observers,
2.5 h of observation each) from 1200 to 1520 AST were made at Adjuntas
on 28 May 1982. At Isabela a total of 20 h of observation (3 observers, 6.7 h
of observation each) was conducted, with a total of 15 h in the morning
(0700-1020 AST), 3 h in the afternoon (1340-1500 AST), and 2 h at night
(1940-2030 AST). Night and afternoon observations were difficult because
of heavy rain during this period (27-29 May 1982). Weevils were present
both at Isabela and Adjuntas and at Isabela they were so numerous that
they had defoliated several papayas near the citrus grove. Larvae were ob-
tained from the U. S. Dept. of Agriculture Laboratory in Mayagiiez. The
source for their laboratory population was the Isabela infestation. At Isabela
and Adjuntas malathion, dicofol, and methomyl were applied once a year
or when pests built up. Insects and spiders were quite abundant at all loca-
tions in Puerto Rico.
Both Isabela and Adjuntas had high humidity and temperatures near
25 C or slightly higher during the afternoon, cooling to no lower than
200C at night. Heavy rains fell in Puerto Rico during the late May observa-
tions, whereas conditions were unusually dry in Florida during summer and
early fall 1981.
Surveys of surface ants were conducted at Forest City on 9 September
1982 and at Isabela on 29 May 1982 with tuna fish placed around flags at 20
stations within the groves. The stations were arranged in two crossing
lines of 10 stations each. After a half hour the baits were checked and ant
samples collected with an aspirator. The ants were then placed in alcohol
and transported to Gainesville, FL for identification.
RESULTS AND DISCUSSION
The dominant predators found in all of the groves were ants (Table 1)
and these accounted for 99% of the predation. The major ant species in-
217
218 Florida Entomologist 66 (2) June, 1983
volved were Tetramorium simillimum Roger, Pheidole dentata Mayr, P.
floridana Emery, and Paratrechina bourbonica (Forel) in Florida (Table 1)
and Pheidole fallax antillensis Forel, T. bicarinatum (Nylander), and Phei-
dole subarmata borenquensis Wheeler in Puerto Rico (Table 2). P. fallax
antillensis was by far the most important of the predators at Isabela, ac-
counting for 88.4% of the 215 predation events observed. The Florida list
varies from that of Whitcomb et al. (1982) only in the order of importance
of a few species and in rates of predation.
The only non-ant predators observed were an earwig, Labidura riparia
(Pallas) (Dermaptera: Labiduridae) and a jumping spider, Corythalia
canosa (Walckenaer) (Araneae: Salticidae). All the Puerto Rican predation
records were for ants. The predation rate was much lower at Lockhart than
at Forest City, with the Puerto Rican groves being closer to Lockhart for
afternoon periods (Forest City 17.1%, Isabela 6.2%, Adjuntas 3.7%, Lock-
hart 2.1%). Whitcomb et al. (1982) found that an average of 47% of the
larvae were consumed by predators at Forest City from 1700 to 0600 EST,
with 62% being consumed in the early morning. At Isabela, Puerto Rico,
8.4% of the larvae (total 2250) that were placed out in the morning were
consumed and 0.0% of the larvae (total 300) placed out at night were
consumed. At night the ground, grass and trees were very wet and droplets
of water were everywhere. The arboreal ant Camponotus ustus Forel was
quite active at night on the trunks of citrus on 27 May, but surface ant ac-
tivity was very low.
Predation by ants in the Florida groves varied from 53% during a 20-min
period at Forest City (15 Sept. 1981, 1430-1450 EST, cloudy and 320C with
60% RH) to 0 at Lockhart on several occasions during the study. This was
most pronounced on 2 Oct. 1981 with 7 20-min observations of 0 under
TABLE 1. MEAN TOTAL PREDATION EVENTS FOR 3 OBSERVATION DAYS OF
Diaprepes abbreviatus NEONATE LARVAE1 AT FOREST CITY AND
LOCKHART, FL, DURING THE AFTERNOON (1200-1530 EST, BETWEEN
31 AUGUST-3 OCTOBER 1981).
Observed predation events per
2 h observation/day
(1200 larvae)
Forest City Lockhart
Predator species X SE X SE
Tetramorium simillimum Roger 69.3 39.9 2.0 1.4
Pheidole floridana Emery 59.7 18.6 4.3 5.4
Pheidole dentata Mayr 58.3 41.0 0.7 0.9
Paratrechina bourbonica (Forel) 5.3 6.8 14.0 0.3
Pheidole moerens Wheeler 3.7 4.5 2.0 2.8
Conomyrma flavopecta (M. R. Smith) 3.7 2.6 0.0 0.0
Labidura riparia (Pallas) 2.0 1.6 0.0 0.0
Pheidole dentigula M. R. Smith 0.7 0.9 0.0 0.0
Brachymyrmex obscurior Forel 1.0 1.4 0.3 0.5
Pheidole morrisi Forel 0.0 0.0 0.3 0.5
Corythalia canosa (Walckenaer) 0.0 0.0 0.3 0.5
Total 203.7 23.9
Richman et al: Predation of Diaprepes Larvae 219
TABLE 2. TOTAL PREDATION EVENTS ON Diaprepes abbreviatus NEONATE
LARVAE FOR MORNING, AFTERNOON AND NIGHT PERIODS AT ISABELA
AND AFTERNOON PERIODS AT ADJUNTAS, PUERTO RICO IN LATE
MAY, 1982. CORRECTED FOR 2 H/DAY (FOR 900 LARVAE).
Isabela Adjuntas
Predator species Morning Afternoon Night' (Afternoon)
Pheidole fallax
antillensis Forel 25.3 27.0 0.0 6.8
Pheidole subarmata
borenquensis Wheeler 0.4 0.0 0.0 2.6
Tetramorium bicarinatum
(Nylander) 0.5 0.0 0.0 2.6
Cardiocondyla nuda (Mayr) 1.3 1.0 0.0 0.0
Camponotus ustus Forel 0.9 0.0 0.0 0.0
Solenopsis geminata (Fab.) 0.0 0.0 0.0 0.0
Hypopenera opaciceps (Mayr) 0.0 0.0 0.0 1.7
Brachymymex obscurior
Forel 0.0 0.0 0.0 0.9
Ochaetomyrmex
auropunctata (Roger) 0.0 0.0 0.0 1.7
Odontomachus bauri Emery 0.0 0.0 0.0 0.9
Cyphomyrmex rimosus
(Spinola) 0.0 0.0 0.0 0.9
Total 28.4 28.0 0.0 18.1
1Very wet in grove. Based on only 120 min observation (3 observers 90 min each).
sunny, warm (270C or higher), and dry (RH ca. 40%) conditions. Although
there was some increase of ant activity toward the late afternoon, differences
in humidity and cloud cover often masked this effect. Predation was
usually higher during cloudy, humid periods than during sunny and dry,
or rainy periods at both Forest City and Lockhart. A chi square analysis of
differences between early and late afternoon periods at Forest City showed
significance at the 0.05 level, with late afternoon having the higher level
of predation. A similar analysis among 20-min observation subperiods within
each of the 1 h periods and among days of observation also showed some
significance, some of the 20-min observation subperiods having significantly
higher predation on the same day and/or on different days. Variation among
time periods and among days was thus quite high. The Puerto Rican data
were not easily compared in this manner.
The major difference between our study at Forest City and that of Whit-
comb et al. (1982) is that during part of the time (afternoon) in which
many of the larvae of D. abbreviatus apparently drop from the trees, the
predation rates were less than 1/2 as high as they were in evening to morn-
ing (Table 3). For the most part, the same species of predators were in-
volved, but these were less active during the afternoon. Because of the
slightly different techniques involved it was not possible to compare these
data statistically.
The earlier observations using plain petri dishes are somewhat com-
parable to the technique employed by Whitcomb et al. (1982), and these
Florida Entomologist 66 (2)
TABLE 3. PERCENT PREDATION OF NEONATE LARVAE OF Diaprepes abbreviatus
AT FOREST CITY, FL, DURING THE AFTERNOON THROUGH THE
MORNING HOURS 1220 TO 1530 (SUMMER AND EARLY FALL 1981)
AND 1700 TO 0600 EST (SUMMER AND EARLY FALL 1979).
Time period
1200-15301 1700-0600'
(% of (% of
Predator species 3600 larvae) ca. 5560 larvae)
Tetramorium simillimum Roger 5.8 9.8
Pheidole floridana Emery 5.0 11.3
Pheidole dentata Mayr 4.9 14.6
Paratrechina bourbonica (Forel) 0.4 3.0
Pheidole moerens Wheeler 0.3 0.9
Conomyrma flavopecta M. R. Smith 0.3 1.8
Labidura riparia (Pallas) 0.2 less than 0.1
Pheidole dentigula M. R. Smith 0.2 0.0
Brachymyrmex obscurior Forel less than 0.1 0.0
Pheidole morrisi Forel 0.0 3.0
Corythalia canosa (Walckenaer) 0.0 less than 0.1
Solenopsis invicta Buren 0.0 2.3
Paratrechina vividula (Nylander) 0.0 0.4
Total 17.1+ 47.1+
1Current study.
2Converted from data in Whitcomb, Gowan and Buren, 1982.
resulted in a 1.1% predation rate at Forest City for 30 July 1981 (1200
larvae, interrupted by rain).
The observations in Puerto Rico did not produce the expected large
number of predation events for night and morning. Because of extreme wet
conditions at night, few ants were seen on the surface of the soil. By mid
to late morning the soil and grass had dried sufficiently for numerous ants
to be active on the soil. A comparison between the soil ants trapped at 20
flagged bait stations at Isabela, Puerto Rico (29 May 1982), and at Forest
City, FL (9 Sept. 1982) is shown in Table 4. Only 2 ant species were shared
(Tetramorium bicarinatum and Brachymyrmex obscurior Forel). However,
both faunas were characterized by Pheidole, Tetramorium, and Solenopsis
species. The Forest City grove contained a very large number of colonies of
Conomyrma flavopecta (M. R. Smith), and 70% of the bait stations at-
tracted this species. The ant was, however, not an efficient predator of
Diaprepes larvae and accounted for only 1.8% of the predation events ob-
served at Forest City in the current study and 3.8% of the predation events
recorded by Whitcomb et al. (1982). At Isabela Pheidole fallax antillensis
and Tetramorium bicarinatum were attracted to equal numbers of bait
stations (65%).
CONCLUSIONS
Since it appears that larvae drop from egg masses in the afternoon and
evening (Jones and Schroeder, in press) the data presented in this report
indicate that the predation of neonate larvae of Diaprepes abbreviatus on
220
June, 1983
Richman et al: Predation of Diaprepes Larvae 221
TABLE 4. ANTS ATTRACTED TO SURFACE TUNA FISH BAIT (IN LATE MORNING-
EARLY AFTERNOON) AT ISABELA, PUERTO RICO, 29 MAY 1982 AND
FOREST CITY FLORIDA, 9 SEPTEMBER 1982.
% of baits visited'
Ant species Isabela Forest City
Pheidole fallax antillensis Forel 65 0
Pheidole subarmata borenquensis Wheeler 30 0
Tetramorium bicarinatum (Nylander) 65 5
Solenopsis geminata (Fab.) 25 0
Brachymyrmex obscurior Forel 15 20
Conomyrma flavopecta (M. R. Smith) 0 70
Pheidole dentata Mayr 0 35
Pheidole floridana Emery 0 30
Tetramorium simillimum Roger 0 10
Solenopsis invicta Buren 0 10
Crematogaster ashmeadi Mayr 0 5
'Sums to more than 100% because baits were often visited by more than 1 species.
the soil surface in Florida may not be as important as previously reported
for late afternoon through morning (Whitcomb et al. 1982). Soil surface
predation was relatively low in the Puerto Rican citrus groves utilized in
this study regardless of the time of day. Recent studies on egg predation
(Richman, Buren, and Whitcomb, unpublished data) and parasitism and
predation on subterranean larvae (Roman and Beavers in press; Beavers,
McCoy and Kaplan unpublished data; Klein and Buren unpublished data)
indicated that eggs and subterranean larvae may be more vulnerable to
attack than freshly dropped larvae on the soil surface. The predators include
various ants and the parasites are primarily nematodes. Weather conditions
may also cause heavy mortality among neonates falling on dry soil (D.
Richman pers. obs.). The egg parasite Tetrastichus haitensis Gehan is ap-
parently not an effective control agent (Schroeder and Beavers 1977).
ACKNOWLEDGEMENTS
We would like to thank Drs. William J. Schroeder, J. B. Beavers, and
Ira Jones (U.S. Dept. of Agric., ARS Plymouth, FL) for their help. We
would also like to thank Drs. Deborah Green and Agripino Perez of the
Isabela Agricultural Research Station, University of Puerto Rico, for their
help with our work in Puerto Rico. Takuji Hayakawa, Anne Trambarulo,
James Trager and Mary K. Elvin of the University of Florida, Dept. of
Entomology and Nematology, helped with the field work or analysis of data.
This study was partially supported by P.L. 89-108, Section 406 Grant
TAD 124914329 entitled "Control of Diaprepes and other root weevils".
Florida Agric. Exp. Sta. Journal Series No. 3797.
REFERENCES CITED
JONES, I. F., AND W. J. SCHROEDER. 1983. Study of first instar Diaprepes
abbreviatus and larval activity for control purposes. J. Econ. Ent.
76(3): (in press).
Florida Entomologist 66 (2)
ROMAN, J., AND J. B. BEAVERS. A survey of Puerto Rican soils for entomo-
genous nematodes which attack Diaprepes abbreviatus (Coleoptera:
Curculionidae). J. Agric. Univ. Puerto Rico (in press).
SCHROEDER, W. J., AND J. B. BEAVERS. 1977. Citrus root weevils in Florida:
Identification, biology and control. Proc. Int. Soc. Citriculture 2: 498-
500.
WETMORE, A. E. 1916. Birds of Puerto Rico. Bull. United States Dept. Agric.
362: 1-140.
WHITCOMB, W. H., T. D. GOWAN, AND W. F. BUREN. 1982. Predators of
Diaprepes abbreviatus (Coleoptera: Curculionidae) larvae. Florida
Ent. 65(1): 150-8.
WOLCOTT, G. N. 1924. The food of Puerto Rican lizard. J. Dept. Agric.
Puerto Rico. 7: 1-37.
WOODRUFF, R. E. 1964. A Puerto Rican weevil new to the United States
(Coleoptera: Curculionidae). Florida Dept. Agric. Div. Plant Ind.
Ent. Circ. 30: 1-2.
-- 1968. The present status of a West Indian weevil (Diaprepes ab-
breviatus (L.)) in Florida (Coleoptera: Curculionidae) Florida Dept.
Agric. Div. Plant Ind. Ent. Circ. 77: 1-4.
VERTICAL AND TEMPORAL ASPECTS OF
ALSYNITE PANEL SAMPLING FOR ADULT
STOMOXYS CALCITRANS (L.) (DIPTERA: MUSCIDAE)
EDWARD F. GERSABECK1 AND RICHARD W. MERRITT
Department of Entomology, Michigan State University,
East Lansing, MI 48824 USA
ABSTRACT
A 45 cm X 3 m vertical Alsynite@ panel coated with Tack Trap@ was
used to study adult flight behavior of Stomoxys calcitrans (L.) at 3 different
land use areas in Michigan. Data indicated 2 daily plateaus of stable fly
activity at 1000 to 1300 h and 1500 to 1800 h. Ninety-five percent of the
total trap catch occurred below 180 cm and between 0800 and 2000 h. More
females than males were trapped closer to the ground. The largest number of
both male and female flies were captured where equine host activity was
greatest.
RESUME
Para estudiar el comportamiento de vuelo de adults de Stomoxys calci-
trans en Michigan en tres areas donde el uso de la tierra era diferente, se
usaron panels verticales de Alsynite de 45 cm x 3 m en tomafio cubiertos
con Tack Trap. Los datos indicaron altiplanos diaries de actividad de S.
calcitrans a las 1000-1300 h y a las 1500-1800 h. Noventa y cinco % de las
moscas que se encontraron en las trampas fueron capturadas debajo de 180
cm y entire 0800 y 2000 h. Mas cerca del suelo mas hembras que machos
fueron capturadas. El numero m6s grande de machos y hembras fue
capturadas debajo de 180 cm y entire 0800 y 2000 h. Mas cerca del suelo
'USDA-APHIS-VS-IP; US Embassy-Tuxtla Gutierrez; P. O. Box 3087, Laredo, TX 78041
USA
222
June, 1983
Gersabeck & Merritt: Sampling Stable Flies 223
mas hembras que machos fueron capturadas. El numero mas grande de
machos y hembras fue capturado donde la actividad equina era mAxima.
Traps constructed of Alsynite@ translucent panels covered with Tack
Trap@ have been used in sampling adult populations of Stomoxys calcitrans
(L.) (Williams 1973, Williams and Rogers 1976, Ruff 1979, Berry et al.
1981, Buschman and Patterson 1981).
In 1976, Williams and Rogers used this trap to examine vertical flight
behavior of stable flies by exposing panel traps for one week intervals at
selected heights below 22.9 m. Ninety-one percent of their total catch oc-
curred when traps were placed at 0.3 and 1.2 m heights above the ground;
the remaining 9% were captured at heights of 2.1, 8.5, 15.2 and 22.9 m.
Typical use of these traps has been to index adult activity at a certain
location. If more precise information is to be obtained from these traps, such
as population parameters based upon mark-release-capture techniques, then
the operational characteristics of these traps need to be more precisely de-
fined. The objectives of this study were to: 1) determine if an optimal loca-
tion for Alsynite@ panels near ground level exists to maximize attraction of
adult stable flies; and 2) determine temporal changes of male and female
stable fly attractancy to Alsynite panels over a 24 h period.
MATERIALS AND METHODS
Location
The study was conducted in the North Central United States on the
island of Mackinac which lies 12 km off the north eastern coast of Michi-
gan's lower peninsula. The island has a surface of ca. 990 ha with ca. 13 km
of shoreline. The vegetation is primarily northern coniferous forest with
ornamental trees and shrubs introduced into populated areas.
The island's economy and recreation have developed around tourism.
During the summer, approximately 500 to 600 horses are brought to the
island and utilized either as saddle horses or to pull carriages and wagons.
The resultant feed and waste from the horses together with garbage from
residents and tourists create a favorable organic media for the development
of S. calcitrans.
Sampling
To test for vertical and temporal activity patterns, 10 translucent
Alsynite@ panels (30 cm x 45 cm) were coated with Tack Trap and ar-
ranged in a continuous vertical column on one stake. Thus, each experi-
mental set of panels formed an Alsynite@ rectangle of 45 cm X 3 m with the
base of the first panel located at ground level. Although other investigators
have used Alsynite@ in a cross configuration, one plane was considered
sufficient to examine stable fly behavior and attractancy in relation to the
panels. Each set of panels was left in place for 1 h. At the end of that hour,
the panels were labeled, removed from the stake, and placed within a
screened enclosure. This enclosure prevented additional flies from attaching
to the panel while in transit to the laboratory. New panels were then placed
on the stake for another hour of exposure.
224 Florida Entomologist 66(2) June, 1983
Twelve experimental runs were conducted, each consisting of 24 se-
quential hours of exposure. In 4 of the 12 runs, 10 panels were changed
every hour for 24 h. In the remaining 8 runs, only one set of 10 panels was
left in place during the time interval 2200 to 0600 h since less than 0.1% of
the total trap catch occurred during this exposure period.
Panels were returned to the laboratory and the following data were
recorded from each panel: 1) the number of female and male stable flies;
2) height interval; 3) date; 4) time of exposure; and 5) site location. In
addition, temperature and humidity were recorded in the field using a hygro-
thermograph.
The experiment was run at 3 locations on the island during the months
of July and August 1980. One site was a dray operation where 8 horses were
stabled. This site was outside the downtown city area and away from main
roads used by animals and people. The second site was within the city area
adjacent to a high-use road; however, no horses were held in corrals or
stables at this site. The last site was a commercial horse drawn carriage
tour operation that maintained ca. 300 horses next to a main route for horse
drawn wagons.
Variability in the raw data required transformation to percent of total
capture at each time or height interval. An analysis of variance was used to
determine if there were significant differences between trap catches at
different heights and times. Comparison among treatment means of sex ratios
over time and at different heights were made using the Duncan's Multiple
Range Test. The probability level of 0.05 was used to denote significant
differences.
RESULTS AND DISCUSSION
Temporal Factors
The mean number of flies caught per panel, the percentage of total trap
catch, female to male sex ratios and associated meteorological factors over
time are presented in Table 1. Stable fly activity, as reflected in trap
catch, increased steadily from 0600 h to 1000 h, where it then remained at a
plateau from 1000 h to 1500 h. A decrease in trap catch followed and a second
plateau occurred between 1500 h and 1800 h (Table 1). This was followed
by a steady decline until 0600 the following day. The 2 plateaus of increased
fly activity occurred when air temperature was increasing or near maximum
and humidity was decreasing or at the minimum daily value. These plateaus
occurred within the temperature range of 21 to 32C during which time
Voegtline and co-workers (1965) also observed heavy biting activity of
stable flies in the upper peninsula of Michigan. In contrast to our findings,
LaBrecque et al. (1975) in Florida reported that flight activity was minimal
during peaks of temperature and light intensity. Other workers have also
reported 2 daily peaks of adult stable fly activity (Hafez and Gamal-Eddins
1959, Kunz and Monty 1976) but at other times of the day, depending on
the study area.
Less than 0.1% of the total trap catch occurred between 2200 and
0600 h (Table 1). This low trap catch reflects both the inability of the
panels to be attractive in the absence of the sunlight and the decrease in
stable fly activity that occurs during nocturnal conditions (Miller et al.
1969).
Gersabeck & Merritt: Sampling Stable Flies
TABLE 1. MEAN NUMBER OF STABLE FLIES CAPTURED FOR EACH HOUR OF
EXPOSURE WITH ASSOCIATED SEX RATIOS AND METEOROLOGICAL
FACTORS (N = 12).
% of the Sex Rel.
X Flies Total Flies Ratio Temp. Humid.
Time Captured' Captured F/M'1 C %
0600-0700 23h 0.9 1.20c
0800 28h 1.1 1.26c 19 92
0900 74g 2.9 1.41b
1000 110f 4.3 1.34b 20 86
1100 179d 7.0 1.39b
1200 174d 6.8 1.59a 22 78
1300 212dc 8.3 1.61a
1400 148e 5.8 1.66a 23 75
1500 169ed 6.6 1.61a
1600 338a 13.2 1.41b 24 71
1700 284b 11.1 1.34b
1800 338a 13.2 1.24c 25 68
1900 253cb 9.9 1.23c
2000 143e 5.6 1.40b 23 82
2100 54g 2.1 1.63a
2200 26h 1.0 1.61a 22 89
2200-0600 5i 0.2 -
'Values followed by different letters differ significantly (p < 0.05).
Female to male ratios for adult flies captured from 0600 to 1000 h and
1600 to 1900 h were significantly lower than sex ratios occurring during
other time intervals (Table 1). These data suggested that perhaps a greater
proportion of male stable flies were actively flying in the early morning
and late afternoon. This hypothesis is supported by the work of Charlwood
and Lopes (1980) who found increased biting activity of male stable flies in
Brazil during similar time periods. Buschman and Patterson (1981) also
found that male stable flies captured on fiber glass panels consistently out-
numbered females and appeared earlier in the day than did females.
Height Factors
Figure 1 shows the percentage of total trap catch as a function of height
above ground level with respective sex ratios for each height. Partitioning
of the trap catch revealed that 95% of the catch occurred below 180 cm. Sex
ratios occurring below and above 90 cmn were significantly different from
each other with more females than males being captured closer to ground
level.
A significant difference occurred between those flies caught below 60 cm
and flies caught above this height. Since stable flies had the opportunity
to land anywhere between 0 and 3 m, these data indicated that optimal trap
placement for maximizing stable fly attraction to Alsynite panels would
occur below a 60 cm height above the ground.
225
Florida Entomologist 66(2)
60 120 180 240
300
Height (cm) Above Surface
Fig. 1. Distribution of trap catch (vertical bars) and sex ratios (open
circles) of stable flies captured at 30 cm intervals above ground level.
Location
Adult stable fly movement and aggregation at a particular site have
generally been associated with mating and host seeking (Gatehouse and
Lewis 1973, Buschman and Patterson 1981) and in the case of females, search
for a suitable ovipositional media. Thus, adult activity at a particular
location should reflect both host activity and the presence of organic wastes.
Table 2 separates panel trap catches and sex ratios by location. The
largest total trap catch and the greatest female to male ratio occurred
where the resident host density was greatest. The lowest female to male
ratio occurred where resident host density was lowest, although the number
of working horses passing both these sites was similar.
Overall sex ratios at all locations for trapped stable flies ranged from
1.50 to 1.61: 1 females to male. These ratios lie within the normal population
range of 1.4 to 1.6: 1 (females to male) reported by Kuntz and Monty (1976)
as determined by sweepnet sampling. Since the sex ratios at these locations
were not significantly different, the observed variation in total trap catch
could not be attributed to changes in activity patterns of a particular sex.
TABLE 2.
MEAN NUMBER OF CAPTURED STABLE FLIES, ASSOCIATED SEX RATIOS,
AND NUMBER OF STABLED HORSES AT EACH STUDY LOCATION (N =
4 REPS/SITE).
Mean no. No. of
captured Mean F/M stabled
Location flies/24th ratio horses
Dray 2223 1.57 : 1 8
City 2543 1.50 : 1 0
Barn 2914 1.61 : 1 300
226
June, 1983
Gersabeck & Merritt: Sampling Stable Flies
SUMMARY AND CONCLUSIONS
This study showed that flight activity of S. calcitrans as reflected by
Alsynite panel catch varies both through time and space. In northern
Michigan the greatest periods of adult fly activity occurred from 1000 to
1300 h and from 1500 to 1800 h. At all times of the day more females than
males were captured in the traps. Over 95% of the total trap catch occurred
between 0800 and 2000 h and below 180 cm.
For those researchers interested in biting fly population estimates based
on mark-release-recapture techniques, this study clearly indicated that
variation in the numbers of flies captured can be reduced by the standardiza-
tion of Alsynite panel trap height and placement (i.e., distance from host
activity).
Research supported by USDA/SEA Competitive Animal Health Grant
No. 59-2261-0-2-060-0 awarded to R. W. Merritt, R. S. Patterson, and I. V.
Berry.
REFERENCES
BERRY, I. L., P. J. SCHOLL, AND J. I. SHUGART. 1981. A mark and recapture
procedure for estimating population sizes of adult stable flies. Environ.
Ent. 10: 88-93.
BUSCHMAN, L. L., AND R. S. PATTERSON. 1981. Assembly, mating, and
thermoregulating behavior of stable flies under field conditions.
Environ. Ent. 10: 16-21.
CHARLWOOD, J. D., AND J. LOPES. 1980. The age structure and biting behavior
of Stomoxys calcitrans (L.) (Diptera:Muscidae) from Manaus,
.Brazil. Bull. Ent. Res. 70: 549-55.
GATEHOUSE, A. G., AND C. T. LEWIS. 1973. Host location behavior of
Stomoxys calcitrans (L.). Ent. Exp. appl. 16: 275-90.
HAFEZ, M., AND F. M. GAMAL-EDDIN. 1959. Ecological studies on Stomoxys
calcitrans (L.) and sitiens Rond. in Egypt with suggestions on their
control. Bull. Soc. Ent. Egypt 63: 245-83.
KUNZ, S. E., AND J. MONTY. 1976. Biology and ecology of Stomoxys nigra
Macquart and Stomoxys calcitrans (L.) (Diptera:Muscidae) in
Mauritius. Bull. Ent. Res. 66: 745-55.
LABRECQUE, G. C., D. L. BAILEY, D. W. MEIFERT, AND D. E. WEIDHAAS. 1975.
Density estimates and daily mortality rate evaluations of stable fly
(Stomoxys calcitrans (L.) (Diptera:Muscidae)) populations in field
cages. Canadian Ent. 107: 597-600.
MILLER, J. A., J. L. ESCHLE, AND I. L. BERRY. 1969. Patterns of flight activity
in livestock insects. 1. Preliminary testing of a system for recording
flight activity of the stable fly. Ann. Ent. Soc. America 62: 1046-50.
RUFF, J. J. 1979. Trapping effectiveness of several combinations of colors
and textures of sticky traps for stable flies, Stomoxys calcitrans (L.).
Mosq. News. 39: 290-2.
VOEGTLINE, A. C., G. W. OZBURN, AND G. 'D. GILL. 1965. The relation of
weather to biting activity of Stomoxys calcitrans (L.) along Lake
Superior. Papers. Michigan Sci. Arts Letters. 1: 107-14.
WILLIAMS, D. F. 1973. Sticky traps for sampling populations of Stomoxys
calcitrans (L.). J. Econ. Ent. 66: 1279-80.
-- AND A. J. ROGERS. 1976. Vertical and lateral distribution of stable
flies in northwestern Florida. J. Med. Ent. 13: 95-8.
227
Florida Entomologist 66 (2)
AUTOGENY IN THE CERATOPOGONIDAE:
LITERATURE AND NOTES
J. R. LINLEY
Florida Medical Entomology Laboratory
University of Florida, Institute of Food and Agricultural Sciences
P. O. Box 520, Vero Beach, FL 32961 USA
ABSTRACT
A brief review is presented, in the form of a species list with appropri-
ate literature citations, of the occurrence of autogeny in the Ceratopogonidae.
There are 38 species in which the phenomenon is now described. Notes on
parthenogenesis, fecundity and ovarian development are appended.
RESUME
Se present un breve repaso, en forma de una lista de species con citas
bibliograficas pertinentes, sobre lo occurrencia de la autogenesis en la Cera-
topogonidae. Existen 38 species en las cuales el fen6meno ha sido ob-
servado. Se afiaden notas adicionales sobre various puntos de interns.
It is commonly observed that the literature tends to grow quietly and
sometimes without being noticed as much as it should. When I recently
finished a list of the autogenous species of Ceratopogonidae, the final com-
pilation of 38 species came as something of a surprise. Since Glukhova and
Dubrovskaya (1972) had earlier mentioned only 20 species (total reported),
it seemed useful to provide an updated review. It has purposely been kept
in the simple form of an alphabetical species list, with appropriate literature
citations (Table 1). Accompanying notes emphasize points of interest or
certain clarifications regarding the identity of species or the evidence for
autogeny when this seemed less than completely conclusive.
The Russian fauna has been surveyed quite extensively, such that
Glukhova and Dubrovskaya (1972) were able to indicate 6 of 21 Russian
species as being autogenous. They also listed the Russian publications re-
ferring to autogeny or anautogeny in these species and the localities in which
the studies were made. Since I have not read all these Russian papers in
the original, I have not copied the citations into Table 1. A translation of
Glukhova and Dubrovskaya (1972) is available (I am grateful to E. T.
Schmidtmann and B. A. Mullens for this translation), to which I refer the
reader. A few citations from the Russian literature are given and these
are papers that have been available in translation for some time. Table 1
is given below and, following it, the notes.
Dasyhelea spp.
Although specific documentation appear to be lacking, autogeny is
thought to be very widespread and perhaps universal in this genus of over
300 species (e.g. Downes 1955).
Culicoides obscuripennis
The report of autogeny in this species is referred to Murphy (1961), but
228
June, 1983
Linley: Autogeny in Ceratopogonidae
TABLE 1. AUTOGENY IN THE CERATOPOGONIDAE: SPECIES AND LITERATURE
CITATIONS.
Literature source
Culicoides alaskensis Wirth
C. austeni Carter, Ingram and Macfie
C. bambusicola Lutz
C. barbosai Wirth and Blanton
belkini (Wirth and Arnaud)
bermudensis Williams
brevipalpis Delfinado
circumscriptus Kieffer
C. dendrophilus Amosova
C. furens (Poey)
C. filicinus Gornostaeva and
Gachegova
C. gigas Root and Hoffman
C. hollensis (Melander and Brues)
C. impunctatus Goetghebuer
C. machardyi Campbell and
Pelham-Clinton
C. mackerrasi (Lee and Reye)
C. marmoratus (Skuse)
C. melleus (Coquillett)
C. mississippiensis Hoffman
C. obscuripennis Clastrier and Wirth
C. praetermissus Carter, Ingram and
Macfie
C. pycnostictus Ingram and Macfie
C. riethi Kieffer
C. salinarius Kieffer
C. segnis Campbell and Pelham-
Clinton
C. subimmaculatus Lee and Reye
C. sanguisuga (Coquillett)
C. waringi (Lee and Reye)
C. variipennis albertensis Wirth
and Jones
Downes (1958)
Murphy (1961)
Lee (1968)
Linley (1965)
Linley (1966)
Duval et al. (1978)
Williams (1961)
A. L. Dyce (pers. comm.)
Glukhova (1958)
Becker (1960)
Boorman and Goddard (1970b)
Glukhova and Dubrovskaya
(1972)
Amosova (1959)
Glukhova and Dubrovskaya
(1972)
Linley (1965)
Linley (1966)
Linley (1969)
Linley et al. (1970)
Koch and Axtell (1978)
Glukhova and Dubrovskaya
(1972)
Downes (1958)
Koch and Axtell (1978)
Boorman and Goddard (1970b)
Glukhova and Dubrovskaya
(1972)
Dyce and Murray (1967)
Kay (1973)
Henry and Adkins (1973)
Linley and Hinds (1976)
Koch and Axtell (1978)
Davis (1981)
Murphy (1961)
Dipeolu and Ogunrinade (1976)
Dipeolu and Ogunrinade (1976)
Glukhova and Dubrovskaya
(1972)
Boorman (1974)
Glukhova and Dubrovskaya
(1972)
Boorman and Goddard (1970a)
Edwards (1982)
Jamnback (1965)
Dyce and Murray (1967)
Downes (1958)
Species
229
Florida Entomologist 66 (2)
TABLE 1. CONTINUED.
Species
Literature source
C. wisconsinensis Jones
Leptoconops (Holoconops) bequaerti
(Kieffer)
L. (H.) kerteszi (Kieffer)
L. (H.) linleyi Wirth and Atchley
Leptoconops (Leptoconops) bezzii (Noe)
L. (L.) carter Hoffman
L. (L.) irritans (Noe)
Forcipomyia inornatipennis (Austen)
F. squamipennis Ingram and Macfie
Dasyhelea spp.
Mullens and Schmidtmann
(1981)
Linley (1968)
Majori et al. (1970)
Rees et al. (1971)
Linley (1982)
Bettini et al. (1969a)
Schmidtmann (1976)
Schmidtmann and Washino
(1982)
Bettini et al. (1969b)
Kaufmann (1974)
Kaufmann (1975)
See notes
Culicoides obscuripennis is not named in Murphy's paper. It is the species
he refers to as breeding in mangrove swamps and distinguishable, by virtue
of its pronounced melanism, from C. austeni. Cornet et al. (1974) have dis-
cussed the identity of species studied by Murphy and have pointed out that
this dark species was redescribed as C. obscuripennis by Clastrier and Wirth
(1961).
C. praetermissus, C. pycnostictus
The evidence for autogeny in these 2 species (Dipeolu and Ogunrinade
1976) seems fairly conclusive. The authors found the ovarian follicles in
various stages of development, from Mer's Stage I to Stage IV, in newly
emerged females of both species. The fat bodies seemed more generously de-
veloped at emergence than in species judged to be anautogenous.
C. segnis
Evidence for autogeny in this species, while inconclusive, is at least
suggestive. Boorman and Goddard (1970a) found that most females taken
in light traps were gravid, yet none was ever seen engorged or with traces
of blood.
AUTOGENY AND PARTHENOGENSIS
In addition to being autogenous, there are 2 species in which partheno-
genesis is confirmed. In Bermuda, Williams (1961) managed to keep 2 virgin
females of C. bermudensis (reared singly from larvae) alive for long
enough to obtain a total of 38 eggs. All of these hatched within 48 h. C.
bermudensis is also suspected of being paedogenetic (Kline and Axtell 1977).
In Colombia, Lee (1968) observed embryonation in eggs produced auto-
genously and laid by virgin C. bambusicola. Becker (1961) suspected, but did
not specifically demonstrate parthenogenesis in C. circumscriptus in Scot-
land.
June, 1983
230
Linley: Autogeny in Ceratopogonidae
PRECOCIOUS DEVELOPMENT OF EGGS
There are a few species in which the condition of the ovaries has been
examined at the time of emergence of autogenous females from the pupa. In
some of these, ovarian development is precocious in the sense that the
follicles are well advanced towards maturity in freshly emerged adults. In
Jamaica, a small form of Leptoconops (H.) becquaerti was autogenous (the
large form from the same larval habitats was not), and 97.5% of follicles in
females dissected within 3 h of emergence were in early Stage IV of develop-
ment (Linley 1968). The closely related species, L. (H.) linleyi was recently
examined (Linley 1982) and is similar. In females less than 1 h old,
68.0% of the follicles were in early Stage IV. Among Culicoides, females of
C. barbosai and C. furens at emergence had ovaries developed to Stages IIB
and III, and IIA and IIB, respectively (Linley 1966). Newly emerged C.
dendrophilus had ovaries in Stages IIA to III (Amosova 1959), and in
C. waringi, ovaries were in Stages IIB or III (Dyce and Murray 1967).
Mullens and Schmidtmann (1981) have recently examined female C. wis-
consinensis within 30 min of emergence and found most follicles already in
late Stage III.
VARIABLE PREVALENCE OF AUTOGENY
Prevalence is a term used to denote differences between localities with
respect to the proportion of the population in which autogeny occurs. A few
species are known to show such variation. In 2 Jamaican populations of C.
barbosai, for example, 97% and 72% of the females produced eggs auto-
genously, while among 6 populations of C. furens, the proportions varied
from 91% to 0% (Linley 1966). The collections were simultaneous and the
collection sites in both instances were only a few miles apart. When the
C. furens population at Vero Beach, Florida, was examined weekly for 13
months, all females from emergence traps produced eggs autogenously
(Linley et al. 1970). C. hollensis from North Carolina has been described as
autogenous by Koch and Axtell (1978), but all individuals from South
Carolina were anautogenous (Henry and Adkins 1973).
SEASONAL VARIATION IN FECUNDITY
C. furens (Linley et al. 1970) and C. melleus (Linley and Hinds 1976)
populations in Florida have been sampled continuously for slightly over a
year in each case to measure seasonal changes in size and fecundity of the
autogenous females. All females collected in both populations throughout the
year produced a first batch of eggs autogenously. The seasonal trends in
size and fecundity were similar, with female size (measured by wing length)
inversely proportional to ambient temperature and fecundity substantially
higher in larger females that emerged in the cooler months. The numbers
of eggs matured per female varied seasonally by about a factor of 2 in the
C. furens population, and 3 in the C. melleus.
MULTIPLE EGG BATCHES PRODUCED AUTOGENOUSLY
Schmidtmann (1976) and, more comprehensively, Schmidtmann and
Washino (1982) have reported evidence indicating that, in some populations
231
Florida Entomologist 66 (2)
June, 1983
of Leptoconops (L.) carter, the females produce not only the first clutch of
eggs autogenously, but also achieve autogenous development of the secondary
follicles (in now parous flies) to early Stage IV. This situation is unique in
Ceratopogonidae within the context of present knowledge. Studies to confirm
and further define the phenomenon would be valuable.
Florida Agricultural Experiment Station Journal Series No. 4075.
REFERENCES CITED
AMOSOVA, I. S. 1959. On the gonotrophic relationships within the genus
Culicoides (Diptera: Heleidae). Ent. Rev. USSR. 38: 774-89.
BECKER, P. 1960. Observations on the feeding and mating of Culicoides
circumscriptus Kieffer (Diptera: Ceratopogonidae). Proc. R. Ent.
Soc. London 35: 6-11.
S1961. Observations on the life cycle and immature stages of
Culicoides circumscriptus Kieff. (Diptera, Ceratopogonidae). Proc.
R. Soc. Edinburgh 67: 363-86.
BETTINI, S., B. MAJOR, E. FINIZIO, AND G. PIERDOMINICI. 1969a. Richerche
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Linley: Autogeny in Ceratopogonidae 233
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MATING BEHAVIOR OF WILD ANASTREPHA
FRATERCULUS (DIPTERA: TEPHRITIDAE) ON A
CAGED HOST TREE
JOAO S. MORGANTE, ALDO MALAVASI
Departamento de Biologia, Instituto de Bioci8ncias,
Universidade de Sio Paulo. Caixa Postal 11.461-S. Paulo
S. Paulo, Brazil
AND
RONALD J. PROKOPY
Department of Entomology, University of Massachusetts,
Amherst, MA 01003, USA
ABSTRACT
The mating behavior of the South American fruit fly, Anastrepha
fraterculus (Wiedemann) is described, following release of wild virgin
flies onto a field-caged host tree (guava). Sexual encounters and attempted
copulations occurred almost exclusively from 0700-0900 hours and were
confined to leaf nodes or the bottom surface of leaves, where males stationed
themselves, often forming leks, and appeared to emit a sex pheromone
which was attractive to virgin females. There were no male visitations or
sexual encounters on green guava fruit, even though females, after mating,
frequently oviposited there.
RESUME
En este trabajo se describe el comportamiento sexual de la mosca de
frutas suramericana, Anastrepha fraterculus (Wiedemann), en el arbol de
guayaba, el cual fue cubierto por una tela para confinar las moscas virgenes.
Los encuentros sexuales y tentativas de copulaci6n ocurrieron casi exclusiva-
mente entire 0700-0900 horas, principalmente en los peciolos en la face in-
^^
__ __
Morgante et al.: Behavior of Anestrepha fraterculus 235
ferior de las hojas donde los machos frecuentemente forman grupos lekss)
que aparentemente emiten feromona sexual la cual atrae las hembras
virgenes. Entretanto, machos y encuentros sexuales no fueron observados
en las guayabas verdes aunque se hayan observado hembras haciendo ovi-
posici6n despues de la c6pula.
Anastrepha fraterculus (Wiedemann) a widely-distributed tephritid
species whose range extends from the Rio Grande Valley in Texas to Argen-
tina and Chile, is a serious pest of many tropical and subtropical fruits
(Malavasi et al. 1980). The damage is caused by the larvae, which feed in
the fleshy parts of fruit rendering it unfit for human consumption.
In several tephritids, visual, auditory, and chemical stimuli are im-
portant in sexual attraction and recognition. With respect to Anastrepha
flies, Nation (1972) described for A. suspense (Loew) the courtship be-
havior and evidence for a male-released sex pheromone that appears to be
released during distention of the abdominal pleural region and anal pouch,
accompanied by rapid wing fanning. It has been identified by Nation (1975)
as a blend of 2 alcohols and 2 lactone esters. Male A. suspense also produce
distinct sounds through wing fanning which appear to be sexually stimulat-
ing to the females (Sharp and Webb 1977, Webb and Sharp 1976).
Studies of the mating behavior of tephritid flies under semi-natural or
natural conditions have been reviewed by Fletcher (1977), Prokopy (1977,
1980), and Burke (1981). Recently, we described the distribution and ac-
tivities of A. fraterculus flies on host and non-host trees in nature (Mala-
vasi et al. 1983). In that study, we observed 26 encounters between males
and females, 2 of which resulted in attempted copulation. Here, we present
evidence suggesting the existence of a sex pheromone in A. fraterculus and
report mating behavior of wild flies released onto a field-caged guava tree,
a major host of this species.
MATERIALS AND METHODS
All of the flies used in all tests emerged from pupae obtained from in-
fested guavas collected in Itaquera, 30 km away from Sio Paulo. Six hours
after eclosion, the sexes were segregated and maintained in laboratory cages
at 250C and 14 h light:10 h darkness. The flies were provided with water and
a mixture of 3 parts sucrose:1 part enzymatic yeast hydrolysate.
To gain information on whether A. fraterculus males release a sex
pheromone, we used an olfactometer similar to that of Kobayashi et al.
(1978). For each of the 21 replicates, 50 virgin females (30-40 days old)
were placed in the large (30 x 30 x 30 cm) fly-holding chamber of the olfacto-
meter. From 0800-1000 h, the females were assayed for response to the test
cage containing either 10 males or no flies (control). The number of females
attracted to each test cage after 2 hours was compared by the paired t sta-
tistical test.
All tests were conducted in a single field cage (3 m diam, 2.7 m tall)
placed over a naturally-growing guava tree, Psidium guajava L. (1.2 m diam,
2.4 m tall), shaded by an overstory of Tibouchina trees. The guava tree had
740 leaves but no naturally growing fruit. Each test day 25 green, freshly-
picked washed guava fruits (2-3 cm diam) were hung by wire from the
guava tree. These green fruit were readily accepted as oviposition sites. We
Florida Entomologist 66 (2)
also hung 4 yellow (ripe), freshly-picked guava fruits (3-4 cm diam),
readily accepted as adult fly feeding sites. Flies (20-30 days old) were
placed in a laboratory room with the natural photoperiod (0600-1900 h)
at 230C three days before release, and were released shortly after dusk
(1800 h) into the cage. Release was accomplished by careful transfer of
each fly from a laboratory cage to the bottom surface of a leaf, using a
small piece of paper under the foretarsi as an aid. Thirty flies of each sex
were released for each of the 5 replicates. The duration of each test was one
day. All of the flies from a previous replicate were removed from the cage
prior to the next replicate. Observations began at dawn (ca. 0600 h) and
ended at dusk (ca. 1800 h). Our presence in the cage had no discernible
effect on the flies.
Once per hour from 0600 to 1800 h, we took a census of the entire tree
and cage wall and ceiling for ca. 15 min, recording the location and activity
of each fly observed. During the interval between each hourly census, we
charted the activities of randomly selected flies for up to 5 min each.
RESULTS
LABORATORY EVIDENCE FOR SEX PHEROMONE: In each test, a mean of 7.8
virgin females were attracted toward a test cage containing males while a
mean of 2.5 virgin females were attracted toward an empty test cage (t =
2.16; P < 0.05).
FIELD CAGE ACTIVITIES:
Location of flies. During the entire day, more males and females, and
considerably more mating pairs, were located on the guava tree than on the
cage wall or ceiling (Table 1). Of those on the tree, the majority of non-
copulating individuals were located on the bottom surface of leaves. Some
females were on the top surface of leaves, green fruit (where ovipositions
were frequent), and ripe fruit, with none on branches or leaf nodes. Some
males were on ripe fruit and leaf nodes, with few or none on other struc-
tures. All males observed on leaf nodes were "calling". All mating pairs
were on the bottom surface of leaves, although copulation may have been
initiated elsewhere.
Location of encounters. The majority of male:female and male:male
encounters, attempted copulations, and successful copulations occurred on
leaf nodes (Table 2). A few male:mating-pair encounters occurred on the
bottom surface of leaves. No female:female or female:mating-pair en-
counters were observed.
Time of copulation initiation. Copulation was initiated almost exclusively
between 0700 and 0900 hours, with very little sexual activity before or after
this period (Table 3).
Courtship behavior. All flies spent the night on the bottom surface of
leaves or on the cage wall or ceiling. Male "calling" began when the
temperature reached ca. 180C and light intensity ca. 800 lux (i.e. at ca. 0700
h each day at summer), and ceased at ca. 1030 h. The predominant sites of
calling coincided with the predominant sites of encounters: leaf nodes and
the bottom surface of leaves. A calling male usually spent ca. 30 min at the
same site before leaving. Calling males appeared to form leks consisting of
3-8 individuals, with each male located on a separate leaf node or leaf and
236
June, 1983
Morgante et al.: Behavior of Anestrepha fraterculus 237
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Morgante et alu Behavior of Anestrepha fraterculus 239
TABLE 3. TIME OF COPULATION INITIATION OF A. fraterculus ON A FIELD-
CAGED GUAVA TREE.
Bottom
Leaf surface Cage
Time Nodes of leaves Branches wall Total
0600-0700 1 0 0 0 1
0700-0800 16 2 1 2 21
0800-0900 6 3 3 1 13
0900-1000 1 1 0 0 2
1000-1800 0 0 0 0 0
all within ca. 50 cm of one another. Leks were particularly pronounced in
the parts of the tree receiving the greatest sunlight. The same was true of
lek formation on a non-host tree in nature (Malavasi et al. 1983).
"Calling" in A. fraterculus was observed to be similar to that described
for A. suspense by Nation (1972). During this behavior the abdominal
pleural region and anal pouch are distended, with both wings at right angles
to the body axis and the coastal edges oriented upward. Periodically, the
wings are vibrated rapidly (fanned) for ca. 5 sec.
The male was the resident in all 36 observed intersexual encounters. In
most of these, the male was in the process of calling when the female
arrived. Typically, the arrival of a female in flight occurred in 2 stages.
First, the female landed within ca. 15 cm of the male, and then walked
toward the stationary male, approaching frontally. The male and female
continued to face each other at a distance of ca. 2 cm, and remained so for
ca. 3 sec. Then the male leaped over the head of the female and onto her
abdomen. The copulating pair immediately proceeded to walk to the bottom
surface of a leaf. Of 29 observed mating pairs, 7 remained in copula for
less than 1 h, 7 for 1-2 h, 12 for 2-3 h, and 3 for more than 3 h. As il-
lustrated by the data in Table 2, calling males encountered and attempted
copulation with other males just as frequently as with females, suggesting
little or no ability of males to discriminate between the sexes prior to
contact.
DISCUSSION
Although female attraction toward males may have been partly on the
basis of male visual or auditory stimuli, we believe male-released sex
pheromone during the process of "calling" was the principal stimulus.
This study shows that copulation in A. fraterculus on a caged host
(guava) tree was initiated almost exclusively on leaf nodes on the bottom
surface of leaves. It was the female which invariably flew toward a some-
what stationary male, rather than the male pursuing a female. Our evidence
strongly suggests that stationary males on leaf nodes emit a sex pheromone
attractive to virgin females. Leaf nodes and the bottom surface of leaves
are the major sites of calling by male A. fraterculus on uncaged trees in
nature (Malavasi et al. 1983). As in other non-temperate tephritids (e.g.
Nation 1972, Feron 1962) the principal strategy employed by an A. suspense
male to attract a female appeared to be a release of sex pheromone coupled
Florida Entomologist 66(2) -
with possible sound produced by wing vibration. The fact that calling and
mating initiation were confined almost exclusively to the hours of 0700-0900
may be important in pre-mating reproductive isolation between different
Anastrepha species simultaneously utilizing the same plant. For example,
Anastrepha ludens (Loew) and A. suspense are known to mate in late
afternoon or early evening (Baker et al. 1944, Perdomo et al. 1975).
No inter or intrasexual encounters among A. fraterculus were observed
on green fruit, even though already-mated females were frequently ob-
served there attempting oviposition. This is in strong contrast to the
situation in Rhagoletis flies, where males often attempt to force-mate al-
ready mated, ovipositing females on fruit (Prokopy 1980). The total ab-
sence of A. fratercules males from green fruit suggests that the oviposition
deterring pheromone deposited by the female on fruit after egglaying
(Prokopy et al. 1982) has no arrestant effect on the males.
This study, together with that of Malavasi et al. (1983), provides be-
havioral-ecological evidence useful in elucidating patterns of divergence
between A. fraterculus and other Anastrepha species.
We thank Carlos Wei, Sergio R. Matioli and Roseli Zanelatto for labora-
tory assistance. This research was supported by Conselho Nacional de De-
senvolvimento Cientifico e Tecnologico, CNPq Brazil, grant number
40.2323/80 (Dr. Prokopy's visit) and number 40.0680/80 (Plano Integrado de
Genetica).
REFERENCES CITED
BAKER, A. C., W. E. STONE, C. C. PLUMMER, AND M. MCPHAIL. 1944. A re-
view of studies on the Mexican fruitfly and related Mexican species.
USDA Misc. Publ. 531, 155 p.
BURK, T. 1981. Signaling and sex in acalypterate flies. Florida Ent. 64:
30-43.
FERON, M. 1962. L'instinct de reproduction Chex la mouche mediterranean
des fruits Ceratitis capitata Wied. (Dipt. Trypetidae). Comporte-
ment sexuel-Comportement de ponte. Rev. Path. Veg. Ent. Agric.
France 41: 1-129.
FLETCHER, B. S. 1977. Behavioral responses of Diptera to pheromones,
allomones, and kairomones. Pages 127-48 In: Chemical Control
of Insect Behavior (Ed.: H. H. Shorey and J. J. Mckelvey) New
York, Wiley.
KOBAYASHI, R. M., K. OHINATA, D. L. CHAMBERS, AND M. S. FUJIMOTO. 1978.
Sex pheromones of the oriental fruit fly and the melon fly: Mating be-
havior, bioassay method, and attraction of females by live males
and by suspected pheromone glands of males. Environ. Ent. 7: 107-12.
MALAVASI, A., J. S. MORGANTE, AND R. A. ZUCCHI. 1980. Biologia de "moscas-
das-frutas" (Diptera: Tephritidae) I. Lista de hospedeiros e dis-
tribuicao geografica. Rev. Brasil. Bio. 40: 9-16.
AND R. J. PROKOPY. 1983. Distribution and activities of
Anastrepha fraterculus flies on host and non-host trees in nature. Ann.
Ent. Soc. America 76: 286-92.
NATION, J. L. 1972. Courtship behavior and evidence for a sex attractant in
the male Caribbean fruit fly, Anastrepha suspense. Ann. Ent. Soc.
America 65: 1364-7.
1975. The sex pheromone blend of Caribbean fruit fly males: Isola-
tion, biological activity, and partial chemical characterization.
Environ. Ent. 4: 27-30.
240
June, 1983
Morgante et al.: Behavior of Anestrepha fraterculus 241
PERDOMO, A. J., R. M. BARANOWSKI, AND J. L. NATION. 1975. Recaptures of
virgin female Caribbean fruit flies from traps baited with males.
Florida Ent. 58: 291-5.
PROKOPY, R. J. 1977. Stimuli influencing trophic relations in Tephritidae.
Colloq. Intern. CNRS 265: 305-36.
1980. Mating behavior of frugivorous Tephritidae in nature. Pages
37-46 In: Proc. Symp. Fruit Fly Problems, XVI Int. Congr. Ent.
Kyoto Nat. Inst. Agric. Sci., Japan.
A. MALAVASI, AND J. S. MORGANTE. 1982. Oviposition deterring
pheromone in Anastrepha fraterculus flies. J. Chem. Ecol. 8: 763-71.
SHARP, J. L., AND J. C. WEBB. 1977. Flight performance and signaling sound
of irradiated or unirradiated Anastrepha suspense. Proc. Hawaiian
Ent. Soc. 22: 525-32.
WEBB, J. C., AND J. L. SHARP. 1976. Analysis and identification of sounds
produced by the male Caribbean fruit ffy, Anastrepha suspense.
American Soc. Agr. Eng. 69: 415-20.
SYSTEMATICS OF ASIOPSOCIDAE (PSOCOPTERA)
INCLUDING PRONOTIOPSOCUS AMAZONICUS
N. GEN. N. SP.
EDWARD L. MOCKFORD
Department of Biological Sciences
Illinois State University
Normal, IL 61761 USA
ABSTRACT
Pronotiopsocus n. gen. is described with P. amazonicus n. sp. as the type
species. The new genus is placed in family Asiopsocidae together with
Asiopsocus Giinther and Notiopsocus Banks. The 3 genera are diagnosed.
Their probable plesiomorphous characters are derived by outgroup com-
parison with superfamily Caecilioidea. These characters are taken to repre-
sent a hypothetical common ancestor of the Asiopsocid genera, which is used
to reconstruct the phylogeny of these genera. Pronotiopsocus represents one
line of the phylogeny with Notiopsocus and Asiopsocus representing a sister
lineage. Geographical distributions suggest great antiquity with a Pangaean
origin of the family. The furcation of the lineage bearing Notiopsocus and
Asiopsocus may have coincided with the initial breakup of Pangaea. Absence
of Notiopsocus from India, Madagascar, and Australia may be due to in-
sufficient sampling of these regions or to the timing of origin and spread of
the genus relative to the breakup of Gondwanaland.
RESUME
Pronotiopsocus gen. n. se describe con el tipo P. amazonicus sp. n. El
nuevo genero se coloca en la familiar Asiopsocidae, junto con Asiopsocus
Giinther y Notiopsocus Banks. Los 3 g6neros se diagnosan. Sus caracteres
plesiomorfos probables se driven por comparaci6n con la superfamilia
Caecilioidea. Estos caracteres se toman como representantes de un ante-
pasado comin hipot6tico de los g6neros asiops6cidos, empleado para la re-
construcci6n de la filogenia de 6stos g6neros. Pronotiopsocus represent una
linea de la filogenia con Notiopsocus y Asiopsocus representando una linea-
Florida Entomologist 66 (2)
hermana. Las distribuciones geogrdficas sugeren gran antiguidad con un
origen Pangean de la familiar. La furcaci6n de una linea llevando Notiopsocus
y Asiopsocus puede haber coincidido con la fragmentaci6n inicial de Pangaea.
La ausencia de Notiopsocus de India, Madagascar, y Australia puede ser
debido a la falta de investigaci6n en 6stas regions, o puede ser debido al
patron temporal del origen y distribuci6n de aquel genero en relaci6n a la
fragmentaci6n de Gonduanalandia.
The family Asiopsocidae Mockford and Garcia Aldrete (1976) was based
on the genera Asiopsocus Giinther and Notiopsocus Banks. It was regarded
by its authors as a phylogenetic sister group with the lineage represented
by families Caeciliidae, Amphipsocidae, Stenopsocidae, and Polypsocidae
(or combinations of these) and was, therefore, regarded as a superfamily
while the latter lineage was also given status as a single superfamily,
Caecilioidea. The 2 superfamilies constitute Group Caecilietae Pearman.
Recently, I have received material from Amazonian Brazil representing
a 3rd genus of Asiopsocidae. The new genus is notable in having among its
characters some identical with Asiopsocus, some identical with Notiopsocus,
some intermediate between these genera and some intermediate between the
set of Asiopsocus plus Notiopsocus and the Superfamily Caecilioidea.
In this paper the new genus and its single included species are described,
the family Asiopsocidae and its genera are defined morphologically, and the
biogeographic and phylogenetic relationships of the genera are discussed.
MATERIALS AND METHODS
The description of Pronotiopsocus amazonicus n. gen., n. sp. is based on 2
female specimens. These are compared with series of 4 species of Notiopsocus
and one of Asiopsocus at hand. All figures were made with aid of a drawing
tube. Measurements were made with a filar micrometer. The micrometer unit
was 0.462 pm. The outer angle of the mandible has not been used previously
to my knowledge. It is taken as follows: assuming that the mandible on a
slide has assumed the flattest possible position, one draws a camera-lucida
outline of the mandible. The 2 mandibles are not mirror images, and I have
consistently used the right mandible in this study, which shows a more
prominent molar lobe (Fig. 2m) and a poorly developed premolar lobe, if
any (Fig. Ip). One draws a line from the most basal point on the median
margin of the molar lobe (Fig. 2b), generally marked by a tiny beard of
spines, to the most basal point on the lateral margin of the mandible (Fig.
21); one constructs a perpendicular to this line on the outline tangent to the
tip of the apical incisor tooth. One draws a line parallel to line bl, crossing
the perpendicular, and tangent to the most distal point on the mandible.
From the point where this line crosses thp perpendicular, one draws a line
to 1. The angle formed by the latter line with bl is the outer angle (Fig. 2a).
A large outer angle, as seen in Caecilioids (range 68-710 for 5 species
representing 5 genera) appears to be correlated with a relatively longer
mandible as compared with the width of its base, while a smaller outer
angle, as seen in Asiopsocids, is generally correlated with a relatively
shorter mandible.
242
June, 1983
Mockford: Systematics of Asiopsocidae
Pronotiopsocus Mockford NEW GENUS
Fig. 1-11
DIAGNOSIS. Male unknown. Female micropterous, wings represented by a
slight bulge on each side of meso- and metatergum. Showing usual characters
of micropterous psocids: ocelli absent, thoracic notal lobes not expressed,
tarsal ctenidia absent, sense cushions of paraprocts greatly reduced.
Antennae broken off and missing beyond scape and pedicel in specimens on
hand. Dorsal ecdysial line of head (Fig. 3) very distinct, with arms reach-
ing nearly to antennal sulci but becoming faint at distal ends. Compound
eyes relatively small but with numerous ommatidia. Labrum ( Fig. 4) with
pigment pattern suggesting broad pair of longitudinal bars. Distal margin of
labrum broad, relatively flat, limited by 2 stylets of moderate length. Distal
inner labral sensilla (Fig. 5) a row consisting from middle outward in either
direction of a, b, c, d (alternating pit-seta-pit-seta); no row of small setae
on outer surface opposite inner sensilla. Mandibles (Fig. 1, 2) with incisor
region relatively elongate; outer angle of mandible = 620. Lacinia (Fig. 6,
6a) with tip broad, with a few broad, rounded denticles of low relief; a
decided bulge on lateral surface before tip. Pretarsal claw (Fig. 7) with
broad, short pulvillus truncated or slightly expanded apically and with long
basal seta. Pearman's organ represented only by 'mirror'. Subgenital plate
(Fig. 8) broadly rounded distally; pigmented area with broad, rounded
median anterior indentation, narrowing and extending to posterior margin
in middle; setae evenly distributed over surface of pigmented area. Ovi-
positor valvulae (Fig. 9): 1st valvula slightly curved, rounded apically;
largely membranous but sclerotized and pigmented basally; 2nd valvula
straight, relatively narrow, rounded apically; 3rd valvula apparently repre-
sented by a large seta and possibly also by a small lobe on posterior margin of
clunium before the seta. Spermatheca (Fig. 10) with broad, rounded,
1 2
,P
Fig. 1-2. Pronotiopsocus amazonicus n.sp. 9. 1.) Left mandible, p = pre-
molar lobe; scale = 0.05 mm. 2.) Right mandible, m = molar lobe; straight
lines and letters b and 1 indicate construction for determining a, the outer
angle of the mandible (see text); scale as in Fig. 1.
243
244 Florida Entomologist 66 (2) June, 1983
slightly sclerotized region around spermapore; duct relatively short, entirely
surrounded by narrow sheath; neck of sack at junction with duct relatively
broad, thickened. Epiproct (Fig. 11) semicircular, bearing several short,
stout setae. Paraproct (Fig. 11) with sense cushion represented by 2 tri-
chobothria with basal florets and a short, stout seta, or only stout seta;
median margin bearing a prominent duplex spine flanked dorsally and
ventrally by a stout seta; numerous other setae confined mostly to medio-
ventral quadrant of paraproct.
TYPE SPECIES. Pronotiopsocus amazonicus n. sp.
Pronotiopsocus amazonicus Mockford NEW SPECIES
Fig. 1-11
Structural features as described for the genus.
MEASUREMENTS (pm). Hind femoral length = 378, hind tibial length =
578, length of 1st hind tarsomere = 180, length of 2nd hind tarsomere = 100.
IO/d (least distance between compound eyes in dorsal view divided by
transverse diameter of compound eye in dorsal view) = 5.29.
COLOR (in alcohol). Compound eyes black. Remainder of head pale tawny
brown, darker on sulci, ecdysial lines, and mandibles, creamy white on areas
of vertex shown unstippled on Fig. 3. Thorax and membranous portion of
abdomen grayish white except sulci, intersegmental lines of thorax, and
slender, transverse dorsal sclerite of base of abdomen tawny brown. Legs
pale tawny brown. Clunium, epiproct, paraprocts, and pigmented area of
subgenital plate pale tawny brown.
HOLOTYPE. Micropterous 9, BRAZIL: Amazonas, Rio Taruma Mirim at
confluence with Rio Negro, approximately 20 km northwest along Rio Negro
from Manaus, 30-IX-1976, J. Adis collector. Paratype, micropterous 9, type
locality, 1-III-1976, J. Adis collector, in tree ecletor traps. The holotype will
be deposited in the Instituto Nacional de Pesquisas da Amazonia, Manaus,
Brazil. The paratype will be deposited for the present in my collection and
ultimately in the Florida Collection of Arthropods, Gainesville, Florida.
DEFINITION OF FAMILY ASIOPSOCIDAE
With the addition of Pronotiopsocus, the Family Asiopsocidae may be
defined by the following characters:
1. lacinial tip broad with denticles of low relief;
2. short labral stylets present;
3. mandible of only moderate length compared to its basal width; outer
mandibular angle 56-620;
4. pulvillus variable: broad, narrow, or absent;
5. winged form with long R-M fusion in hindwing;
6. Pearman's organ represented only by 'mirror' (Pronotiopsocus) or
field of minute spines (Asiopsocus), or absent (Notiopsocus);
7. first and second ovipositor valvulae, when present and distinct, with
distal ends rounded;
8. ventral abdominal vesicles absent (present in most Caecilioidea);
9. apophyses of subgenital plate absent (present in many Caeciliodea).
Mock ford: Systematics of Asiopsocidae 245
8
S \ \ *..._ /
ELt
c-N- /^ CJ ;
'! ^iV V
^"^ C' c"^' ""***-* -'*)
iI
12
:~~--- ILi:~, ,....~
I I----
10
h-------------I
Fig. 8-11. Pronotiopsocus amazonicus n.sp. 9. 3.) Head in dorsal view;
scale = 0.1 mm. 4.) Labruin showing pigmentation pattern; scale = 0.1 mm.
5.) Distal margin of labrum showing inner sensilla (a, b, c, d); scale = 0.1
mm. 6.) Lacinia, scale = 0.5 mm. Fig. 6-a. Lacinial tip; scale = 0.01 mm. 7.)
Pretarsal claw; scale = 0.01 mm. 8.) Subgenital plate; scale = 0.1 mm. 9.)
Ovipositor valvulae; scale = 0.05 mm. 10.) Spermatheca; scale of Fig. 9. 11.)
i ----
246 Florida Entomologist 66 (2) June, 1983
STRUCTURAL COMPARISONS AND PHYLOGENY OF THE
ASIOPSOCID GENERA
The family Asiopsocidae consists at present of the 3 genera Asiopsocus,
Notiopsocus, and Pronotiopsocus. The genera are differentially diagnosed
by the characters in Table 1.
For derivation of a phylogeny of the Asiopsocidae, characters of an
ancestral asiopsocid were synthesized by comparison between each of the
Asiopsocid genera and the Caecilioidea. Structural features of Caecilioidea
were reviewed by Mockford and Garcia Aldrete (1976) and Mockford
(1978). Similarities are taken to indicate plesiomorphous character states
for the Asiopsocidae. Presumed plesiomorphous states which are apomor-
phous in at least one of the Asiopsocid genera are valuable in determining
phylogeny and are discussed below.
Pronotiopsocus resembles the Caecilioidea in possession of a broad pul-
villus, lack of microtrichs of the base of the pretarsal claw, and possession
of the mirror of the coxal organ. Notiopsocus resembles the Caecilioidea in
possession of the mesepisternal sulcus, possession of a sizable sclerotized 3rd
valvula, and in ciliation of the female forewing. Asiopsocus resembles
certain Caecilioids, probably near the base of that group, in its very broad,
markedly denticulate lacinial tip. Both Notiopsocus and Pronotiopsocus re-
semble Caecilioidea in possession of a sclerotized, free 2nd ovipositor val-
vula. The 2nd valvula of Asiopsocus is a membranous bulge on the 3rd or is
lacking entirely. Macroptery, found in most Caecilioidea, and in females of
Notiopsocus and males of Asiopsocus, is presumed to be the ancestral
condition of Asiopsocidae.
Assuming that the 9 states of the above paragraph characterized the
ancestral Asiopsocid, how were the modern genera derived from it? Each
possible pair of these genera shares at least one apomorphous state of the
hypothesized ancestral characters. Therefore, it seems reasonable to try to
adduce a phylogeny in which 2 of the genera are derived as one lineage
and the 3rd as another lineage. Obviously, some parallel changes would
occur in such a phylogenetic pattern. I have chosen the phylogeny contain-
ing the fewest parallel changes, apparently representing the simplest hypo-
thesis. Pronotiopsocus is derived as one lineage and Notiopsocus and
Asiopsocus are derived as a sister line. This pattern contains 3 parallel
changes, while the other possible patterns contain at least 4. The Pronotiopso-
cus lineage is characterized by the apomorphous states of the lacinial tip
being somewhat narrowed, the mesepisternal sulcus being absent, the 3rd
valvula being greatly reduced, and the female being micropterous. The
hypothesized common ancestor of Notiopsocus and Asiopsocus is character-
ized by the apomorphous states of the pulvillus being slender, and the base
of the pretarsal claw bearing microtrichs. The Notiopsocus line derived from
this ancestor is characterized by the apomorphous states of the lacinial tip
being narrowed, the pulvillus being absent, and the male being micropterous.
The Asiopsocus line is characterized by the apomorphous states of the mese-
pisternal sulcus being absent, the 2nd valvula being reduced and mem-
branous, the 3rd valvula being membranous, the female being micropterous,
and wing setae (in the male) being absent.
Mockford: Systematics of Asiopsocidae
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247
Florida Entomologist 66 (2)
GEOGRAPHIC DISTRIBUTION OF ASIOPSOCIDAE
The family Asiopsocidae is known from North America, Eurasia, tropical
South America, and tropical Africa. It is not known from Madagascar,
India, and Australia. Asiopsocus presents a truly bizarre distribution, with
one species in Mongolia (Ginther 1968, 1974), one in Arizona and adjacent
Sonora (Mockford and Garcia Aldrete 1976, Mockford 1977), and one in
Spain (Lienhard 1981). Notiopsocus is entirely neotropical and tropical
African except for one species in southern Florida (personal observation),
and Pronotiopsocus is known only from Amazonian Brazil.
The distribution of Asiopsocus suggests great antiquity. Its relationship
to Notiopsocus, adduced above, suggests origin as the northern vicariant in
the phylogenetic event which led to fragmentation of the common ancestor
of those 2 genera. If this event coincided with the breakup of Pangaea, one
can presumably place it in time as Late Triassic to Early Jurassic, 203-192
m.y.BP (Seyfert and Sirkin 1979). The common ancestor of these 2 genera
must have separated from Pronotiopsocus before that time.
The absence of Asiopsocidae in Madagascar, India, and Australia, though
possibly due to inadequate sampling, may be due to the timing of separation
of plates. If Notiopsocus was spreading eastward from a, let us suppose,
South American region of Origin and reached Africa before that continent's
separation from South America (Late Jurassic to Early Cretaceous, 150-
130 m.y.BP according to Seyfert and Sirkin 1979), it may not have been
able to reach the plate containing Antarctica, Australia, and India, which
separated from that containing Africa and Madagaccar between Late Tri-
assic and Early Jurassic, 205-160 m.y.BP (Seyfert and Sirkin 1979). It also
may not have been able to reach Madagascar, which separated from Africa
between Middle Jurassic and Late Cretaceous, 172-90 m.y.BP (Seyfert and
Sirkin 1979).
Presence of Notiopsocus in Mexico and southern Florida is probably ex-
plainable as part of a post-Pleistocene invasion of tropical South American
insect species into southern North America (Halffter 1964).
ACKNOWLEDGEMENTS
I wish to thank Dr. J. Adis for sending the specimens of Pronotiopsocus
amazonicus and 2 unknown reviewers for several stimulating comments.
REFERENCES CITED
GUNTHER, K. K. 1968. Staublause (Psocoptera) aus der Mongolei. Mitt.
Zool. Mus. Berl. 44: 125-41.
1974. Psocoptera of the Mongolian People's Republic (In Russian
with German summary). Insects of Mongolia 2: 34-50.
HALFFTER, G. 1964. La entomofauna americana. Ideas acerca de su origen y
distribuci6n. Folia Ent. Mex. 6: 1-108.
LIENHARD, C. 1981. Neue und interessante Psocopteren aus Greichenland,
Spanien, und Portugal. Dtsch. Ent. Z. 28: 147-63.
MOCKFORD, E. L. 1977. Asiopsocus sonorensis (Psocoptera:Asiopsocidae) : A
new record, augmented description, and notes on reproductive biology.
Southwest. Nat. 22: 21-9.
1978. A generic classification of family Amphipsocidae (Psocoptera:
Caecilietae). Trans. American Ent. Soc. 107: 249-98.
June, 1983
248
Mockford: Systematics of Asiopsocidae
--- AND A. N. GARCfA ALDRETE. 1976. A new species and notes on the
taxonomic position of Asiopsocus Giinther (Psocoptera). Southwest.
Nat. 21: 335-46.
SEYFERT, C. K., AND L. E. SIRKIN. 1979. Earth history and plate tectonics,
ed. 2. Harper and Row, New York, VIII + 600 p.
FEMALE CALLING BEHAVIOR IN EPHESTIA ELUTELLA
AND E. FIGULILELLA (LEPIDOPTERA: PYRALIDAE)
S. B. KRASNOFF1, K. W. VICK2 AND R. W. MANKIN
Insect Attractants, Behavior, and Basic Biology Research Laboratory,
Agric. Res. Serv., USDA, Gainesville, FL 32604 USA
ABSTRACT
The sexual calling behavior of Ephestia elutella (Hiibner) and E.
figulilella Gregson females was observed in the laboratory under a 14L:10D
photoperiodic regime. E. elutella females showed a broad mid-scotophase
peak of calling activity with low levels of calling continuing throughout the
photophase. E. figulilella females showed a narrower mid-scotophase calling
peak with little or no photophase activity.
RESUME
El comportamiento de las hembras en 'llamar", eso es, atraer a los
machos, fue observado en 2 species, Esphestia elutella (Hubner) y E.
figulilella Gregson en un laboratorio debajo un regimen de fotoperiodo de
14L: 10 D. Hembras de E. elutella mostraron un pico ancho de actividad de
llamar en el medio del period de oscuridad, con niveles bajos de esta activi-
dad siguiendo por todo el period de luz. Las hembras de E. figulilella
mostraron un pico mbs angosto en el medio del period de oscuridad, con
poca o ninguna actividad en el period de luz.
Ephestia elutella (Hiibner) and E. figulilella Gregson frequently co-
occur, and are often found in sympatry with 3 other stored-product infesting
phycitid moths: Plodia interpunctella (Hiibner), E. kuenhiella Zeller, and
E. cautella (Walker) (Reed and I ivingstone 1937, Donahoe et al. 1949,
Hoppe and Levinson 1979). Tnasm-n,, as the primary component of the fe-
male sex pheromone (Z,E)-9,12-ietradecadien-l-ol acetate (ZETA) is the
same in all 5 species (Brady and Daley 1972), and the males of any 1
species can be attracted to and attempt to mate with a female of any of the
other species (Ganyard and Brady 1972, Krasnoff, unpublished data) it was
hypothesized that differences in diel periodicity of sexual activity might
help to maintain reproductive isolation in this group.
Daily rhythms underlying various aspects of sexual behavior and
physiology have been studied in 3 stored-product infesting phycitid moths,
P. interpunctella (Nordlund and Brady 1974, Coffelt et al. 1978), E.
kuenhiella (Calvert and Corbet 1973), and E. cautella (Coffelt et al. 1978).
'Present address: Department of Entomology, Cornell University, Ithaca, NY 14853.
2To whom reprint requests should be directed.
249
250 Florida Entomologist 66 (2) June, 1983
However, the literature contains only anecdotal references to the diel sexual
rhythms of the 2 other stored-products Phycitinae, E. figulilella (Cox 1974),
and E. elutella (Reed and Livingstone 1937).
A laboratory study of the daily calling cycles of E. elutella and E.
figulilella was undertaken to complete the picture of temporal partitioning of
sexual calling in the stored-products Phycitinae, as well as to provide a
foundation for a study of the courtship behavior of these 2 species.
MATERIALS AND METHODS
REARING AND HANDLING. Ephestia elutella were obtained from a colony
maintained since 1961 at the USDA Tobacco Storage Insects Research Unit
in Richmond, VA. E. figulilella were derived from a 15-year-old colony
maintained at the USDA Stored-Product Insects Research Laboratory,
Fresno, CA. Larvae of both species were reared according to the methods
of Silhacek and Miller (1972). An autoclaved diet (Coffelt et al. 1979) was
used for E. figulilella larvae which were found to be highly susceptible to
disease. Cultures of larvae were maintained on a 14L:10D photoperiod at
26 1C and ca. 50% RH. A fluorescent lamp provided ca. 190 lux during
the photophase. Scotophase illumination was ca. 5 lux from a red incandescent
lamp (both measured 1 m from source). Insects were segregated by sex as
pupae.
As female pupae became available they were divided into 3 groups and
placed in 19.5 x 13.5 x 10.0-cm vented-plastic containers. These containers,
in turn were placed in each of three 60 x 60 x 60-cm plexiglass chambers
equipped with opaque cloth hoods so that the photoperiod could be main-
tained without interference from extraneous light. The photocycle in each
chamber was offset from that in each of the others by 8 h. Thus hourly
observations for 24 h of the daily cycle could be taken in 8 h. A light-board
with an array of nine 7.5-W incandescent bulbs (5 white and 4 red) pro-
vided ca. 50 and 10 lux at 30 cm during the photophase and scotophase, re-
spectively. Only red light was used during the scotophase. Insects were
placed in these chambers early in the pupal stadium so that they would
entrain to the offset photocycles before tclosion (Nordlund and Brady 1974).
Temperature in the chambers was 24-27 C.
Insects that emerged each day were segregated into groups of 5-15 in
cylindrical plastic containers (7 cm x 9 cm diam) and provided with a
piece of fluted construction paper as a perch. Hourly counts were made of
the number of calling females ("strong calling" sensu Nordlund and Brady
1974).
In another experiment, conducted only with E. elutella, 3- to 4-day-old
unmated females (ca. 120) were released from vented mason jars placed in
the center of the floor of a 2.5 x 1.4 x 2.0-m room during the last hour of the
photophase. After 15 min a count was taken of the number of females on
each surface (walls, ceiling, floor) and observations were made on the
direction in which each female on the wall was facing (up, down, sideways).
These counts were taken again 4 h later and the results compared with those
of the 1st count.
RESULTS
Both E. elutella and E. figulilella females assume the characteristic phyci-
Krasnoff et al.: Calling Behavior in Ephestia
tine calling posture (Richards and Thomson 1932). The abdomen is flexed
dorsally and the ovipositor is exserted. The female pheromone gland in both
of these species is located on the ventral aspect of the membrane between
the 8th and 9th abdominal segments, and is homologous to that of P. inter-
punctella (Smithwick and Brady 1977a,b).
Although rhythmic retraction and extrusion of the ovipositor has been
noted in several lepidopteran species (Conner 1979 and references therein),
this behavior was not observed in either E. figulilella or E. elutella. Females
of both species, however, beat their antennae while calling (ca. 175 beats/
min for E. figulilella, and 80 beats/min for E. elutella). Often E. elutella
females initiating a calling bout were observed to pulse the dorsally flexed
abdomen vigorously up and down for a few seconds before bringing it into
the full calling position.
The calling behavior of both E. elutella and E. figulilella females showed
distinct diel periodicities. E. elutella females had a broad peak of calling
activity during the scotophase and continued to call at low levels during the
photophase (Fig. 1). E. figulilella females also showed a maximum calling
response during mid-scotophase, but with little if any calling during the
photophase. The breadth of the peak in E. elutella was greater than in E.
figulilella, such that >50%. of E. elutella females exhibited calling from the
2nd through the 8th h of the scotophase, whereas >50% of the E. figulilella
females exhibited calling from only the 3rd through the 6th h.
At 15 minutes after the onset of darkness, most of the females on the
walls were facing up. This simply reflects the fact that they were released
from the floor, flew up to the walls, landed face-up, and remained facing
up until calling began. By the middle of the calling period (4 h after dark)
::.- EPHESTIA ELUTELLA
EPHESTIA FIGULILELLA
.-J
,-j
95
_5
85
65
5
15
wIlL J L3 L4 D4 L5 I DS L6 D6 L7 D7 LB D81 L9
0 10 24 34 48 58 72 82 96 106 120130144154 168178 192
HOURS AFTER ONSET OF FIRST SCOTOPHASE
Fig. 1. Temporal distribution of calling in Ephestia figulilella and E. elutella
females. Percentages are based on at least 25 observations.
Florida Entomologist 66 (2)
most of the females on the walls were facing down (x" = 69.0, p <.005).
An apparent preference for vertical surfaces was detected in E. elutella in
the room-release study in which a significantly higher number of females
(p <.005) took positions on the walls than on the ceiling or floor (Table 1).
DISCUSSION
The periodicity of the calling behavior of E. elutella females in a light:
dark regime resembles that of P. interpunctella (Nordlund and Brady 1974)
in 3 ways: 1) the activity peak is broad, 2) the peak occurs during mid-
scotophase, and 3) moderate levels of calling activity are maintained during
the photophase. These species are, in effect, photoperiodic "generalists",
adapted to living in enclosed stores and food-processing plants cut off from
natural illumination, and subjected to irregular or non-existent photocycles.
In contrast, E. figulilella, an habitu6 of open storage areas, has a sharper
calling peak and lacks daylight calling.
If the information from the present study is integrated with that from
studies of the other species in this complex (Traynier 1970, Steele 1970,
Nordlund and Brady 1974), it may be possible to draw some conclusions
about temporal partitioning of sexual calling in this group (Fig. 2). P.
interpunctella and E. elutella would be likely candidates for temporal
isolation from one another because they often co-occur, and they share not
only the same primary component in the female sex pheromone but a second-
ary component as well (Krasnoff, Vick and Coffelt, unpubl. data). However,
they clearly are not temporally isolated on the basis of female calling
periodicity.
The calling periods of the 3 remaining species do seem to be discrete
enough from each other to possibly provide some degree of reproductive
isolation, assuming that time of day of male responsiveness approximately
corresponds to time of day. of female calling. A comparative study of
naturally occurring populations of these species is needed to determine
whether the temporal position of any species where it co-occurs with
another (or others) in the group, differs from its position where it is isolated
from the otherss. Documentation of this type of character displacement
TABLE 1. LOCATION AND POSITION PREFERENCE FOR Ephestia elutella FE-
MALES IN A ROOM-RELEASE STUDY (FLYING FEMALES WERE NOT
SCORED).
Surface (position)
Wall
(facing (facing (facing
Floor Ceiling up) side) down) x2a
Lights-out
+ 15 min 14 2 82 5 5 10.9
Lights-out
+ 4 h 6 2 27 7 58 15.3
aRelative proportion of total available surface area for each surface (wall, ceiling, floor)
was used to calculate expected values.
252
June, 1983
Krasnoff et al.: Calling Behavior in Ephestia
>- IEphes,/o figulella
I- Ephest/a cautell/ E s f e// Epheshe kuehriel/a
> Ephestia eluella Plodia interpuncle//lla
I I
-j
L-j
-j
dusk dawn
TIME OF DAY
Fig. 2. Diel periodicity of female calling activity in stored-product
Phycitinae.
would support the hypothesis that species-specificity of the diel sexual
activity period actually evolved in a context of reproductive isolation.
REFERENCES CITED
BRADY, U. E., AND R. C. DALEY. 1972. Identification of a sex pheromone
from the raisin moth, Cadra figulilella. Ann. Ent. Soc. America 65:
1356-8.
CALVERT, I., AND S. A. CORBET. 1973. Reproductive maturation and phero-
mone release in the flour moth Anagasta kuehniella (Zeller). J. Ent.
(A) 47: 201-9.
COFFELT, J. A., L. L. SOWER, AND K. W. VICK. 1978. Quantitative analysis
of identified compounds in pheromone gland rinses of Plodia inter-
punctella and Ephestia cautella at different times of day. Environ.
Ent. 7: 502-5.
-- K. W. VICK, L. L. SOWER, AND W. T. MCCLELLAN. 1979. Sex phero-
mone mediated behavior of the navel orangeworm, Amyelois transi-
tella. Environ. Ent. 8: 587-90.
CONNER, W. E. 1979. Chemical attraction and seduction: The courtship of
Utetheisa ornatrix (Lepidoptera, Arctiidae). Ph.D. Dissertation,
Cornell University, Ithaca, New York.
Cox, P. D. 1974. The influence of temperature and humidity on the life-cycles
of Ephestia figulilella Gregson and Ephestia calidella (Guene6) (Lepi-
doptera, Phycitidae). J. Stored Prod. Res. 10: 43-56.
DONOHUE, H. C., P. SIMMONS, D. F. BARNES, G. H. KALOOSTIAN, AND C. K.
FISHER. 1949. Biology of the raisin moth. United States Dept. Agric.
Tech. Bull. 994, 23 p.
GANYARD, M. C., AND U. E. BRADY. 1972. Interspecific attraction in Lepi-
doptera in the field. Ann. Ent. Soc. America 65: 1279-82.
HOPPE, T., AND H. Z. LEVINSON. 1979. Befallserkennung und Population-
suberwachung vorratsschadlicher Motten (phycitinae) in einer Scho-
253
254 Florida Entomologist 66 (2) June, 1983
koladenfabrik mit Hilfe pheromonbekoderter Klubefallen. Anz.
Schadlingskde. Pflanz. Umweltschutz 52: 117.
NORDLUND, D. A., AND U. E. BRADY. 1974. The calling behavior of female
Plodia interpunctella (Hiibner) under two light regimes. Environ.
Ent. 3: 793-6.
REED, W. D., AND E. M. LIVINGSTONE. 1937. Biology of the tobacco moth and
its control in closed storage. Circ. United States Dept. Agric. 422.
RICHARDS, O. W., AND W. S. THOMSON. 1932. A contribution to the study
of the general Ephestia Gn. (including strymax, dyar) and Plodia
larvae. Trans. Ent. Soc. (London) 80: 169-251.
SILHACEK, D. L., AND G. L. MILLER. 1972. Growth and development of the
Indian meal moth, Plodia interpunctella (Lepidoptera: Phycitidae).
Ann. Ent. Soc. America 65: 466-8.
SMITHWICK, E. B., AND U. E. BRADY. 1977a. Site and production of sex
pheromone in developing female Indian meal moths, Plodia inter-
punctella. J. Georgia Ent. Soc. 12: 1-13.
--- AND 1977b. Histology of the sex pheromone gland in develop-
ing female Indian meal moths, Plodia interpunctella. J. Georgia Ent.
Soc. 12: 13-139.
STEELE, R. W. 1970. Copulation and oviposition behavior of Ephestia
cautella (Walker). J. Stored Prod. Res. 6: 429-45.
TRAYNIER, R. M. M. 1970. Sexual behavior of the Mediterranean flour moth
Anagasta kuehniella: Some influences of age, photoperiod and light
intensity. Canadian Ent. 102: 534-40.
NEW CAVERNICOLOUS PROPTOMAPHAGINUS FROM
HISPANIOLA AND MEXICO (COLEOPTERA:
LEIODIDAE: CHOLEVINAE)
STEWART B. PECK
Department of Biology, Carleton University
Ottawa, Ontario KlS 5B6, Canada
ABSTRACT
Two new species of the genus Proptomaphaginus are described: P. his-
paniolensis, collected at an ultraviolet light trap in a bat-guano cave in the
Dominican Republic and at other cave and forest sites on the Caribbean
island of Hispaniola; and P. reddelli from a cave in Oaxaca, Mexico. The
species are not cave-specialized. The evolution of the 6 known species in the
genus is diagrammed and discussed.
RESUME
Se described 2 species nuevas del genero Proptomaphaginus: P. his-
paniolensis, colectada en una trampa de lhz ultraviolet cerca de una cueva
con murci6lagos y guano en la Republica Dominicana y en otros sitios en
cuevas y en selvas en la isla caribe de Hispanola; y P. reddelli de una cueva
en Oaxoca, Mexico. Las species no son especializadas para cuevas. La evo-
luci6n de las 6 species conocidas en el g6nero se discute y se represent por
diagrama.
Proptomaphaginus Szymczakowski contains 3 species from Puerto Rico
Peck: New Cavernicolous Proptomaphaginus
and Cuba, and one in Mexico. This genus has not been found in Jamaica de-
spite extensive searching (Peck 1977). This paper describes 2 new species;
one from the Dominican Republic and Haiti on the Caribbean island of
Hispaniola, and another from Mexico. All these species are probably scaveng-
ers in deep soils or litter of forests or in caves, and none are cave-specialized
in structure.
Proptomaphaginus shares with its sister group, the diverse genus
Ptomaphaginus Portevin of the Indo-Malayan region, the derived characters
of a narrow mesothoracic epimeron, and a comb of equal spines on the outer
face of the fore-tibia. These characters occur in no other New World members
of the tribe Ptomaphagini. Proptomaphaginus is considered to be derived
from a group that originated in the New World tropics and spread to the
Old World tropics of Asia (Szymczakowski 1969, Peck 1981) perhaps in the
Cretaceous. The group survives in the New World only as relicts in soil and
litter (and caves which are environmentally similar) in Mexico and some
of the Greater Antillean islands.
Proptomaphaginus hispaniolensis PECK NEW SPECIES
(Fig. 1-4)
DESCRIPTION: The species matches the detailed description of P. apo-
demus Szymczakowski (1969) of Cuba except in the following characters:
mm 4
Fig. 1-7. Genital structures of Proptomaphaginus. Fig. 1-4. P. hispanio-
lensis. 1) Dorsal view of aedeagus. 2) Right lateral view of aedeagus. 3)
Ventral view of aedeagus tip. 4) Spermatheca. Fig. 5-7. P. reddelli. 5) Dorsal
view of aedeagus. 6) Right lateral view of aedeagus tip. 7) Spermatheca. All
to the same scale.
255
256 Florida Entomologist 66 (2) June, 1983
Length 1.75-2.13 mm. Width: 0.80-0.90 mm. Eyes large, width to head
margin 3.5 times distance from anterior eye margin to antennal insertion.
Antennae not noticeably thin or elongated; reaching base of pronotum when
laid back. Elytra finely striolate, apices truncate in both sexes and
slightly concave. Flight wings fully developed and functional. Mesosternal
carina low and uniformly rounded. Legs normal, not noticeably elongate.
Aedeagus (Fig. 1) uniformly stout and thick, narrowing to apex, with
prominent subequal lateral lobes at tip; apex (Fig. 2) downturned, with 2
lateral swellings, each bearing 2 setae (Fig. 3); ventral blade of tegmen
collar-shaped, with median opening; internal sac (flagellum) long, thin,
twisted; parameres (Fig. 2, 3) shorter than aedeagus, fused to aedeagus,
bearing 2 setae at tip, and 3 others widely spaced before tip. Spermatheca
(Fig. 4) gently curved and progressively swollen to anterior end.
DIAGNOSIS: The characters of the aedeagus distinguish the species from
all other known Proptomaphaginus. The species is most similar to P.
puertoricensis Peck but is distinguished from this species by a more reflexed
and trilobed median lobe of the aedeagal tip.
HOLOTYPE MALE: Dominican Republic. La Romana Province, in cave at
mouth of Rio Chavon, 8-V-78, R. E. Woodruff and G. B. Fairchild, at black-
light near bat guano (in Florida State Collection of Arthropods, FSCA).
Paratypes: with same data; in FSCA (16), Canadian National Collection
(4), Museum of Comparative Zoology (4), and author's collection (6).
OTHER TENTATIVELY ASSIGNED MATERIAL: DOMINICAN REPUBLIC. Con-
stanza, 3-4000 ft., VIII-1938, P. J. Darlington, 1 female (in MCZ). HAITI.
Dept. du Sud. Grotte Counoubois, 1 mi. SSW Camp Perrin, 290 m, 2-XI-79,
J. R. Holsinger, J. H. Stock, et al., 1 S, 2 9 (in author's collection). Grotte
"Ca-August", between Cayes and Jacmel, 15-XI-79, J. Notenboom and L.
Botosaneanu, 4 9 (in Museum of Zoology, University of Amsterdam).
ETYMOLOGY. The name refers to the island of Hispaniola, where the
species occurs.
BIOLOGICAL OBSERVATIONS: The large eyes and fully functional flight
wings indicate that the species is not cave restricted, (i.e., it is a troglophile).
The available material indicates that the species is widely distributed on
Hispaniola (the cave locations are at nearly opposite ends of the island on
the south coast) and is also in montane forests. This is the same distribu-
tional-habitat eurytopy as for P. puertoricensis (Peck 1970). The Haitian
locality of Grotte Counoubois is a large cave, and is one of several described
by Dunn, Schmidt and Taylor (1959). I know of no published summary of
caves in the Dominican Republic.
The method of collecting the large type series (R. E. Woodruff, pers.
comm.) is noteworthy and should be attempted in other tropical guano
caves. A small 6 volt battery-operated blacklight was run over a white
enamel pan in the cave, near a large pile of bat guano. The beetles, as well as
flies of various families, flew into the light and fell into the pan. Many insects
in cave guano piles are attracted to lights. I have collected flies of many
families from guano piles in caves in Jamaica with carbide, gasoline, and
electric light sources (Peck 1975).
Proptomaphaginus reddelli PECK NEW SECIES
(Fig. 5-7)
Peck: New Cavernicolous Proptomaphaginus
DESCRIPTION: The species matches the description of P. microps Peck
(1973) of Mexico except in the following characters: Length 1.75-1.95 mm.
Width 0.73-0.78 mm. Eyes reduced to cluster of about 20 facets, width to
head margin 0.7 times distance from anterior eye margin to antennal in-
sertion. Antennae not noticeably thin or elongate, reaching to base of
pronotum when laid back. Elytra truncate at apex, sutural angles rounded
in both sexes. Flight wings fully developed (presumed to be functional).
Mesosternal carina low and rounded. Aedeagus (Fig. 5) tubular, elongate;
constricted at middle; lateral lobes at tip unequal, left lobe higher and folded
over right lobe (Fig. 6); ventral blade of tegmen thin and elongate; flagel-
lum of internal sac very long, thin, curved; genital orifice shifted to aedeagal
dorsal surface; parameres (Fig. 6) thin, fused to aedeagus, setae visible
only at tip. Spermatheca (Fig. 7) curved, progressively swollen to anterior
end.
DIAGNOSIS: The species is distinguished from P. microps by the aedeagus,
which is more constricted in the middle, and which has a higher left apical
lobe enclosing the genital orifice.
HOLOTYPE MALE: MEXICO. Oaxaca, Cueva de la Finca, 10 km SW Acatlan,
31-XII-76, J. Reddell, D. McKenzie, and A. Grubbs (in Canadian National
Collection). Paratypes: One 8 and 2 9 with same data in author's collection
and one female in Texas Memorial Museum, Austin.
ETYMOLOGY: The species is named for Mr. James Reddell in recognition
of his explorations and other efforts in elucidating the cave fauna of
Mexico.
DIscussION: The species is judged to be normally an occupant of deep
forest litter but one which can also establish populations in caves. This
species, as well as P. microps, and Ptomaphaginus chapman Peck of
Borneo, all have the unusual combination of very reduced eyes but functional
flight wings. Most litter-dwelling beetles are observed to experience wing
reduction and loss of flight ability before eye reduction.
Proptomaphaginus microps Peck, 1973
The species is known from only 2 caves in the Mexican State of San Luis
Potosi. A new cave record follows: MEXICO. San Luis Potosi, Xilitla Plateau,
Gruta de los Muertos, 28-III-80, Dale Pate, 1 & (in author's collection).
Proptomaphaginus apodemus Szymczakowski, 1969
The species is reported only from 15 caves (usually with bat guano)
distributed throughout Cuba in the provinces of Oriente, Las Villas, Cama-
guey, La Habana, Matanzas, and one cave in Pinar del Rio (Decou 1973). A
new cave record follows: CUBA. Pinar del Rio, Cueva Perfecto, 26-XI-81, P.
Beron, 4 S, 3 Y (in collection of National Museum of Bulgaria, Sofia, and
of author).
EVOLUTION AND BIOGEOGRAPHY
An analysis of this monophyletic group has been performed by standard
Hennigian cladistic methods. The hypotheses of character transformation
series (Table 1) were made by out-group comparison with known characters
257
Florida Entomologist 66 (2)
June, 1983
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Peck: New Cavernicolous Proptomaphaginus 259
and character states in other genera of Ptomaphagini. The analysis suggests
a cladogram of the evolution of the genus as in Figure 8. It indicates that
the Mexican species form a group distinct from those of the Antilles, and
that within the Antillian species the 2 species of Cuba are a sister group to
those of Puerto Rico and Hispaniola. The Mexican species possess more
plesiomorphies, and thus may be most like the generic ancestor. The synapo-
morphies of the Antillian species indicate their monophyly, and that they
are not descendants from multiple Antillian occupation by separate Mexican
ancestors. The separation of the Mexican and Antillean stocks would
probably have been in the early Tertiary according to the evolutionary and
biogeographic models of Rosen (1975, 1978).
ACKNOWLEDGMENTS
The collectors and curators of the new material are thanked for making
it available to me for study, especially John Holsinger, R. E. Woodruff, and
J. Reddell. A. F. Newton and anonymous reviewers read the manuscript
and are thanked for their suggested improvements. The figures were inked
by Hana Kukal and Jennifer Read. The abstract was translated by Maria
Jesus Gines. My research on the distribution and evolution of scavenging
leiodid beetles of forests and caves is supported by operating grants from
the Natural Sciences and Engineering Research Council of Canada.
8 P. hisoaniolensis Peck, Hispaniola
6 P. puertoricensis Peck, Puerto Rico
3
2 7 P. darlingtoni (Jeannel, Cuba
P. apodemus Szymczakowski, Cuba
g P. microps Peck, Mexico
5 P. reddelli Peck, Mexico
Fig. 8. Phylogenetic hypothesis of evolution in Proptomaphaginus.
Numbers refer to characters in Table 1. Only development or appearance
of apotypic character states is shown. No synapomorphy has been found
for the pair of Cuban species (P. darlingtoni is known only from the holo-
type). Thus, the species pair from Hispaniola and Puerto Rico could possibly
be a sister group of either of the Cuban species.
REFERENCES CITED
DECOU, V. GH. 1973. Recherches sur les Col6opteres hypog6s de Cuba. III.
Catopidae-Catopinae: Proptomaphaginus apodemus Szymczakowski.
Pages 367-72 In: R6sultats des expeditions biosp6ologiques Cubano-
Roumaines A Cuba. Editura Academiei Republicii Socialiste Romania,
Bucarest, 424 p.
DUNN, R., V. SCHMIDT, AND E. TAYLOR. 1969. A reconnaissance of caves in
Haiti. Speleo-Digest 1959: 300-15.
PECK, S. B. 1970. The Catopinae (Coleoptera: Leiodidae) of Puerto Rico.
Florida Entomologist 66 (2)
Psyche 77. 237-42.
.1973. A review of the cavernicolous Catopinae (Coleoptera:
Leiodidae) of Mexico, Belize, and Guatemala. Assn. Mexican Cave
Studies Bull. 5: 97-106.
-- 1975. The invertebrate fauna of tropical American caves, Part III:
Jamaica, an introduction. Int. J. Speleol. 7: 303-26.
-- 1977. New records and species of Leiodinae and Catopinae (Coleop-
tera: Leiodidae) from Jamaica and Puerto Rico with a discussion of
wing dimorphism. Psyche 83: 243-54.
-- 1981. A new cave-inhabiting Ptomaphaginus beetle from Sarawak
(Leiodidae: Cholevinae). Syst. Ent. 6: 221-4.
Rosen, D. E. 1975. A vicariance model of Caribbean biogeography. Syst.
Zool. 24: 431-64.
1978. Vicariant patterns and historical explanation in biogeography.
Syst. Zool. 27: 159-88.
SZYMCZAKOWSKI, W. 1969. D6couverte d'un representant des Ptomaphagini
A Cuba (avec une esquisse de la syst6matique et la g6on6mie de cette
tribu) (Coleoptera, Catopidae). Acta Zool. Cracoviensia 14: 87-97.
GROWTH PATTERN ALTERATIONS IN FALL
ARMYWORM, SPODOPTERA FRUGIPERDA', LARVAE
AFTER PARASITIZATION BY APANTELES
MARGINIVENTRIS-, CAMPOLETIS GRIOTI", CHELONUS
INSULARIS', AND EIPHOSOMA VITTICOLE3
TOM R. ASHLEY
Insect Attractants, Behavior and Basic Biology Research Laboratory,
Agricultural Research Service, USDA, Gainesville, FL 32604 USA
ABSTRACT
Parasitization of fall armyworm, Spodoptera frugiperda (J. E. Smith),
larvae by Apanteles marginiventris (Cresson), Campoletis grioti
(Blanchard), Chelonus insularis Cresson, and Eiphosoma vitticole Cresson
reduced maximum larval weights by 97, 96, 70, and 62%, respectively com-
pared to 6th instar nonparasitized larvae. Parasitized larvae produced sig-
nificantly less frass and had smaller head capsule widths. Parasitization in-
creased the duration of the instar during which the parasite destroyed its
host. Apanteles marginiventris and C. grioti destroyed host larvae in the
4th instar and larvae parasitized by E. vitticole died in the 5th instar. Eggs
parasitized by C. insularis were destroyed as larvae in the 4th (41%) and
5th (59%) instars. Larvae parasitized by A. marginiventris gained the least
amount of weight, produced the least amount of frass, had the shortest life
expectancy, did not live past the 4th instar, and had the smallest head capsule
widths.
RESUME
El parasitismo de las larvas de Spodoptera frugiperda (J. E. Smith) por
'Lepidoptera: Noctuidae.
2Hymenoptera: Braconidae.
"Hymenoptera: Ichneumonidae.
260
June, 1983
Ashley: Effects of Parasitization
Apanteles marginiventris (Cresson), Campoletis grioti (Blanchard), Chelo-
nus insularis Cresson y Eiphosoma vitticole Cresson redujeron los pesos
miximos de las larvas por 97, 96, 70 y 62%, respectivamente, en comparisi6n
con las larvas del sexto instar no infestadas por parasitos. Las larvas para-
sitadas produjeron menos material fecal y tuvieron las capsulas de las
cabezas mas pequefios. El parasitismo aument6 la duraci6n del instar durante
lo cual el parasito destruy6 su hospedera. Apanteles marginiventris y C.
grioti destruyeron las larvas de las hospederas en el cuarto instar y las larvas
parasitadas por E. vitticole se murieron en el quinto instar. Los individuos
parasitados en el estadio del huevo por C. insularis fueron destruidos como
larvas en el quarto instar (41%) y en el quinto (59%) instar. Las larvas
parasitadas por A. marginiventris ganaron el menos peso, produjeron menos
cantidad de material fecal, tuvieron el indice de longevidad mas corto, no
vivieron mAs que el cuarto instar, y tuvieron menos anchura de la cApsula
de la cabeza.
Parasites of the fall armyworm (FAW), Spodoptera frugiperda (J. E.
Smith), occur throughout North, Central, and South America (Ashley 1979).
The significant reduction in FAW larval populations caused by some of
these parasites (Ashley et al. 1982) establishes the importance of under-
standing any additional effects these natural enemies have on FAW larvae.
If, for example, parasitization affects larval growth rates then the determina-
tion of the age distribution of FAW field populations either by larval size
or head capsule widths must consider parasitization levels for correct de-
velopment of demographic models. In addition, economic thresholds for
FAW larval populations would have to include levels of parasitization pro-
vided parasitization significantly affects food consumption.
The physiological effects of parasitization and their influence on the
host are reviewed by Vinson and Iwantsch (1980). Modifications of host
growth and development such as reductions in size and weight, alterations
in instar duration, and time of pupation as well as modifications in food con-
sumption rates have been reported by Jones et al. (1981), Hegazi et al.
(1978), Lewis (1970), Rahman (1970), Vinson (1972) and others. Although
parasitization has been studied in many hosts no information is available for
parasitization effects on FAW larvae.
The present research identifies some of the principal effects of parasitiza-
tion by Apanteles marginiventris (Cresson), Campoletis grioti (Blanchard),
Chelonus insularis (= texanus) Cresson (Marsh 1978), and Eiphosoma vitti-
cole Cresson on FAW larvae. Chelonus insularis is a solitary egg-larval endo-
parasite and the remaining 3 species are solitary larval endoparasites. The
larval parasites primarily attack first and second instars. Biological data
for A. marginiventris, C. grioti, and C. insularis are found in Boling and
Pitre (1970), Kunnalaca and Mueller (1979), Morey (1971), Pierce and
Hollaway (1912), and Wilson (1933). Equivalent data for E. vitticole are
not available; however, this parasite has b6en recovered from FAW larvae
(Hynes 1941, Labrador 1967).
MATERIALS AND METHODS
Behavioral differences between parasite species necessitated the use of
different methods and containers for mating and oviposition. Circular plastic
containers (7 x 10 cm diam.) having 2 screened vents (1.5 x 3.0 cm) and
Florida Entomologist 66 (2)
containing ca. 50 first instar larvae and 4 cubes (1.5 cm) of FAW diet
(Leppla et al. 1978) were utilized as ovipositional units for A. margini-
ventris and C. grioti. Two pairs of adult parasites less than 4-h old were
placed into each of 3 containers for both species. Adults of C. insularis were
placed in a 24 cm3 plexiglass cage for 2 days prior to introducing FAW eggs,
and after a 24-h-host-exposure period these eggs were transferred to con-
tainers identical to those used for A. marginiventris.
Three pairs of E. vitticole were held in a plexiglass cage (24 cm3) for 3
days prior to the introduction of hosts. A disk of diet (2 x 8 cm diam) was
supported in the center of the cage by 2 pieces of hardware cloth (8 x 5 cm
high) arranged in an "X" configuration. Depressions (0.5 cm deep) were
made over the upper surface of the disk using 0.64 cm hardware cloth be-
cause the female parasites preferred to search for hosts by probing into
these depressions with their ovipositors. Approximately 100 newly hatched
larvae were placed on the disk and allowed to enter these depressions before
the disk was placed in the ovipositional cage.
The host exposure period for the larval parasites lasted 48 h and all
parasites were continuously supplied with undiluted honey. Larvae emerg-
ing from eggs parasitized by C. insularis remained together for 48 h in
containers like those used for oviposition by A. marginiventris. When the
FAW larvae were 48 h old (all were 2nd instars) they were individually
placed into 30 ml plastic cups containing ca 15 ml of diet. These cups were
sealed with paper lids and every 24 h each larva was transferred into a fresh
diet cup. During transfer the larva's head capsule width, the presence or
absence of an exuvium, and larval and frass weights were recorded. This
procedure continued until a larva pupated or was destroyed by a parasite.
Parasites and FAW larvae were kept at 28+10C, 60 +1.5% RH, and under
a photoperiod of 14:10 LD with a fluorescent light intensity of 800 ft-c. The
analysis is based on 28, 25, 27, and 32 larvae parasitized by A. margini-
ventris, C. grioti, C. insularis, and E. vitticole, respectively and 40 non-
parasitized larvae.
Adults of A. marginiventris came from a laboratory colony started in
1976 from FAW larval collections made at Hastings, Florida. Campoletzs
grioti was imported from Uruguay where it was found parasitizing FAW
larvae in corn (Buckingham, per. comm.). Chelonus insularis was imported
into the United States in 1978 from Bolivia where it was parasitizing FAW
larvae in corn (Mitchell, per. comm.).
An attempt was made to estimate the amount of diet consumed during
a 24 h interval by correcting for weight loss in the diet cups (30 ml) due to
evaporation. The correction factor was derived from diet cups void of larvae
but held for 24 h under the same experimental conditions. Regression
analyses to predict the amount of diet consumed by larvae of a specific age
indicated that a linear model provided optimum prediction and that ad-
ditional precision was not achieved by quadratic or cubic equations. How-
ever, the location of larval feeding activity on the diet (top, side, or within)
coupled with larval metabolism, size, and frass production introduced sub-
stantial error in determining the amount of diet eaten. Therefore, estimates
of diet consumed are not presented.
RESULTS AND DISCUSSION
Parasitization significantly (1% level) reduced daily FAW larval
262
June, 1983
Ashley: Effects of Parasitization 263
weights after the 6th day compared to nonparasitized larvae (Fig. 1A). No
meaningful patterns of weight differences occurred between FAW larvae
parasitized by A. marginiventris and C. grioti. Larvae parasitized by C.
insularis and E. vitticole achieved significantly greater weights (1%
level) compared to larvae parasitized by the other 2 parasites. However, sig-
nificant weight differences were present between larvae parasitized by C.
insularis and E. vitticole on days 10 and 11. Only 28% of the larvae para-
sitized by C. insularis lived past the 9th day and these larvae displayed an
unusual increase in weight prior to destruction by the parasite. Parasitization
by A. mnarginiventris, C. grioti, C. insularis, and E. vitticole reduced maxi-
mum larval weights by 97, 96, 70, and 62%, respectively compared to non-
parasitized 6th instars. The maximum weights achieved for each larva during
each instar were used to determine mean maximum weights (Table 1A).
Parasitization by A. marginiventris and C. grioti, C. insularis and E. vitti-
cole reduced mean maximum weights by 64, 51, 33, and 17%, respectively
compared to nonparasitized 4th instar larvae. Nonparasitized larvae in-
creased their average weight by ca. 265% between successive instars whereas
125
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Fig. 1. Mean comparisons between nonparasitized fall armyworm (FAW)
larvae to FAW larvae parasitized by A. marginiventris (A.m.), C. grioti
(C.g.), C. insularis (C.i.) and E. vitticole (E.v.) for A) larval weights, B)
frass produced, C) head capsule widths, and D) mean instar. Values
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Florida Entomologist 66 (2)
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Ashley: Effects of Parasitization
larvae parasitized by A. marginiventris exhibited the smallest weight gain
of ca. 88% between instars.
Frass production from parasitized and nonparasitized increased with
age until 3 or 4 days prior to the end of the larval stage (Fig. Ib). The most
pronounced decrease occurred in nonparasitized larvae where a reduction
of 106 mg occurred between days 10 and 14. With the exception of hosts
containing immature A. marginiventris, none of the parasitized larvae pro-
duced frass during the last 24 h of their lives. Frass production from all
larvae substantially decreased during the transition period between instars.
Campoletis grioti consumed the entire contents of the host during the last
24 h prior to closing from the FAW larva. After eclosion the parasite larva
formed a cocoon to which the host's cuticle and head capsule were attached.
A mature A. marginiventris larva bored through the side of the host larva
and formed a cocoon on the substrate. The host larva then moved away from
the site of cocoon formation and died within the next 12 to 24 h. Eiphosoma
vitticole and C. insularis caused the host larva to bore into the diet to pupate
early. Frass production in FAW larvae parasitized by these 2 parasites
mimicked that of nonparasitized larvae which enter a gut cleaning stage
prior to pupation.
Parasitization reduced the rate of head capsule growth (Fig. Ic). This
reduction was most pronounced in FAW larvae parasitized by A. margini-
ventris and C. grioti and least pronounced in hosts parasitized by the other
2 parasite species. The apparent reduction in head capsule widths of larvae
parasitized by E. vitticole on day 12 was the result of the FAW larvae
with the larger head capsule widths being destroyed by the parasite on the
preceding day and does not reflect an anatomical reduction in head capsule
widths. Larvae parasitized by C. grioti exhibited the greatest reduction in
the rate of head-capsule-width growth prior to the time of parasite
emergence. A comparison of the mean minimum and mean maximum head
capsule widths for each instar indicated that substantial differences were
not present until the parasitized larva reached the instar during which it
was destroyed by the parasite (Table 1B). This condition was most notice-
able for 4th instar hosts parasitized by A. marginiventris and C. grioti. Al-
though data on larval lengths were not gathered there was a definite re-
duction in the larval length of parasitized larvae.
The rate of advancement from one instar to the next was substantially
reduced in parasitized hosts (Fig. Id). This reduction was most evident in
hosts parasitized by C. grioti and least evident in larvae parasitized by C.
insularis and E. vitticole. Larvae parasitized by A. marginiventris and C.
grioti died as 4th instars and larvae parasitized by E. vitticole died as 5th
instars. Of those larvae parasitized by C. insularis, 41% died in the 4th
instar and 59% died in the 5th instar. Parasitization tended to extend the
duration of the instar during which the parasite destroyed its host rather
than increasing the time spent in earlier ipstars (Table 1C). This trend was
particularly pronounced in larvae parasitized by C. grioti where 4th instars
were present over a 6-day period whereas 4th instars of nonparasitized larvae
were present for 3 days. In addition, both A. marginiventris and C. grioti
spent ca. 44% of their larval lives as 4th instars.
The quantitative differences in larval weights, frass produced, head
capsule widths and mean instars plus noted differences in larval lengths
between parasitized and nonparasitized larvae of the same age demonstrated
265
Florida Entomologist 66 (2)
the significant effects of parasitization on FAW larvae. If parasitization
produced similar effects in field populations of FAW larvae then it may be
necessary to account for the level of parasitization as well as the parasite
species present in establishing economic thresholds and in determining the
correct age distribution of larval populations.
Mention of a commercial or proprietary product does not constitute an
endorsement by USDA.
REFERENCES CITED
ASHLEY, T. R. 1979. Classification and distribution of fall armyworm para-
sites. Florida Ent. 62: 144-53.
V. H. WADDILL, E. R. MITCHELL, AND J. RYE. 1982. Impact of native
parasites on the fall armyworm, Spodoptera frugiperda (Lepidoptera:
Noctuidae), in South Florida and the release of the exotic parasite,
Eiphosoma vitticole (Hymenoptera: Ichnuemonidae). Environ. Ent.
11: 833-7.
BOLING, J. C., AND H. N. PITRE. 1970. Life history of Apanteles margini-
ventris with descriptions of immature stages. J. Kansas Ent. Soc.
43: 465-70.
HEGAZI, E. M., A. M. EL-MINSHAWY, AND S. M. HAMMAD. 1978. Effect
parasitism on digestion and development of Spodoptera littoralis
(Boisd.) larvae. Z. Ang. Ent. 86: 80-5.
HYNES, H. B. N. 1941. Lepidopterous pests of maize in Trinidad. Tropic.
Agric. 19: 194-202.
JONES, D., G. JONES, AND B. D. HAMMOCK. 1981. Developmental and be-
havioural responses of larval Trichoplusia ni to parasitization by an
imported braconid parasite Chelonus sp. Physiol. Ent. 6: 387-94.
KUNNALACA, S., AND A. J. MUELLER. 1979. A laboratory study of Apanteles
marginiventris, a parasite of green cloverworm. Environ. Ent. 8:
365-8.
LABRADOR, J. R. 1967. Estudios de Biologia y Combate del Gusano Cogollero
del Maiz Laphygma frugiperda (S. & A.). Maracaibo, Universidad
del Zulia, Facultad de Agronomia. 83 p.
LEPPLA, N. C., P. V. VAIL, AND J. R. RYE. 1982. Mass rearing the cabbage
looper. In: E. G. King and N. C. Leppla, [eds.], Advances and
Challenges in Insect Rearing. USDA Tech. Bull. (In press).
LEWIS, W. J. 1970. Study of species and instars of larval Heliothis parasi-
tized by Micropletis croceipes. J. Econ. Ent. 63: 363-5.
MARSH, P. M. 1978. The braconid parasites (Hymenoptera) of Heliothis
species (Lepidoptera: Noctuidae). Proc. Ent. Soc. Washington 80:
15-36.
MOREY, C. S. 1971. Biologia de Campoletis grioti (Blanchard) (Hymen.:
Ichneumonidae) Parasito de la "Lagaria Cogollera del Maiz" Spodop-
tera frugiperda (J. E. Smith). Revista Peruana Ent. 14: 263-71.
PIERCE, W. D., AND T. E. HOLLOWAY. 1912. Notes on the biology of Chelonus
texanus. J. Econ. Ent. 5: 425-8.
RAHMAN, M. 1970. Effect of parasitization on food consumption of Pieris
rapae larvae. J. Econ. Ent. 63: 820-1.
VINSON, S. B. 1972. Effect of the parasitoid, Campoletis sororensis on the
growth of its host, Heliothis virescens. J. Insect Physiol. 18: 1509-14.
AND G. F. IWANTSCH. 1980. Host regulation by insect parasitoids.
Quart. Rev. Biol. 55: 143-65.
WILSON, J. W. 1933. The biology of parasites and predators of Laphygma
exigua Huebner reared during the season of 1932. Florida Ent. 17:
1-15.
266
June, 19833
Ashley et al.: Parasitization of Fall Armyworm Larvae 267
PARASITIZATION OF FALL ARMYWORM LARVAE
ON VOLUNTEER CORN, BERMUDAGRASS,
AND PARAGRASS
T. R. ASHLEY, C. S. BARFIELD', V. H. WADDILL2, AND E. R. MITCHELL
Insect Attractants, Behavior, and Basic Biology Research Laboratory
Agricultural Research Service, USDA Gainesville, FL 32604 USA
ABSTRACT
Fall armyworm larvae, Spodoptera frugiperda (J. E. Smith), were
collected from volunteer corn at Homestead, FL from April through August
and from Bermudagrass, Cynondon dactylon (L.), and paragrass, Brachiarie
mutica (L.), in August and September at Belle Glade, FL. Chelonus insularis
Cresson, Temelucha sp., and Apanteles marginiventris (Cresson) parasitized
44, 11, and 5% of all larvae collected, respectively. The principal parasite in
volunteer corn was C. insularis and A. marginiventris was primarily found
in Bermudagrass. Larvae collected from corn were the most frequently at-
tacked with 78 and 72% of the 3rd and 4th instars parasitized, respectively.
Larval abundance and parasitization rates in corn were greatest during the
months of June and July.
RESUME
Se colectaron las larvas de Sodoptera frugiperda (J. E. Smith) de maiz
voluntario en Homestead, FL. desde Abril hasta Agosto y de past de
Bermuda, Cynodon dactylon (L.) y de Brachiarie mutica (L.) en agosto y
septiembre en Belle Glade, FL. Chelonus insularis Cresson, Temelucha sp., y
Apanteles marginiventris (Cresson) parasitaron 44, 11, y 5% de todas las
larvas colectadas, respectivamente. El parasite principal de S. frugiperda
en el maiz fue C. insularis; A. marginiventris fue encontrado en S. frugi-
perda primariamente en past de Bermuda. Las larvas colectadas del maiz
fueron atacadas mas frequentamente, con 78 y 72% de las larvas del tercer
y del cuarto instar respectivamente parasitadas. La abundancia de larvas y
los percentages del parasitismo de S. frugiperda en maiz fueron maximos
en los meses de Junio y Julio.
The fall armyworm (FAW), Spodoptera frugiperda (J. E. Smith), is a
polyphagous, mobile pest that annually attacks corn and other crops, es-
pecially gramineous species, throughout the southeastern United States.
Fall armyworm populations can be found during the entire year in South
Florida and Texas; it is commonly believed that this pest migrates from
these areas into uninfested regions farther north each spring (Mitchell
1979).
Physical environment, available host plants, and natural mortality factors
may act singly or in combination to explain the annual distribution patterns
and densities of FAW populations (Barfield and Stimac 1981). Natural
mortality inflicted on FAW larvae by parasites in both agricultural and
wild host-plant communities is believed to play a substantial role in density
1Department of Entomology and Nematology, University of Florida, Gainesville FL
32611 USA.
2Agricultural Research and Education Center, University of Florida, Homestead, FL 33030
USA.
Florida Entomologist 66 (2)
regulation (Barfield et al. 1980). Studies analyzing larval populations in
southern Florida have been restricted to cultivated corn (Ashley et al.
1980, 1982). The study described here was conducted to identify parasites
attacking FAW larvae on volunteer corn, Bermudagrass, Cynondon
dactylon (L.) and paragrass, Brachiarie mutica, (L.). Each of these plant
species constitutes a potentially significant host for FAW populations and
associated parasites throughout much of the year in South Florida.
MATERIAL AND METHODS
FAW larvae were collected weekly from volunteer corn, commencing in
April 1979 and continuing through August 1979. The corn plants were
located within a 5 m radius of FAW pheromone traps. The sampling pro-
cedure consisted of harvesting several corn plants near each trap. Fall army-
worm larvae were removed from the plants in the laboratory and placed
into individual 30-ml plastic cups containing diet. The head capsule width
of each larva was recorded when it was placed initially in the cup. Larvae
were held in these cups until each larva's fate was determined. Collections
in Bermudagrass occurred on 21 August 1979 and 17 September 1979 and in
Paragrass on 17 September 1979 in Belle Glade, FL; all collections were
made with sweep nets. Both grasses were growing adjacent to sugarcane
fields and no pheromone traps were present in the collecting areas.
Two methods were used to calculate percent parasitization. The first
method utilized all FAW larvae in a particular category regardless of their
size or age. The second method used only larvae with head capsule widths
within the range of head capsule widths parasitized by a particular parasite.
The latter method eliminated larvae that were too large or old (as indicated
by head capsule widths) for parasitization by a particular parasite. Thus, a
more precise determination of the parasite's impact on larvae that it can
successfully parasitize can be made.
RESULTS AND DISCUSSION
Numbers of FAW larvae collected from corn, Bermudagrass and Para-
grass were 1453, 162, and 33 and parasitization rates were 66, 54, and 22%,
respectively. Parasitization levels for all FAW larvae regardless of host
plant indicated that Chelonus insularis Cresson had a rate 4 times greater
than the next most prevalent parasite Temelucha sp. (Table 1). Apanteles
marginiventris (Cresson) was the third most abundant parasite followed
by the tachinids and the remaining hymenopterous parasites, Rogas laphy-
gmae Viereck, Meteorus autographae Muesebeck, and Ophion sp. Each
parasite species emerged from larvae having a particular range of head
capsule widths (Table 1) and these head capsule widths were indicative of
both larval age and size. Percentages for parasitization were also calculated
based on the number of available larvae. A larva was considered available
for parasitization if its head capsule width was within the range of head
capsule widths parasitized by a particular parasite. Slight increases in para-
sitization levels for most of the parasites occurred when this method was
used with the most dramatic increase occurring in Ophion sp. Discounting
time of collection, parasitization rates for C. insularis and Temelucha sp.
were substantially higher in corn than in either of the 2 grasses. Apanteles
268
June, 1983
Ashley et al.: Parasitization of Fall Armyworm Larvae 269
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Ashley et al.: Parasitization of Fall Armyworm Larvae 271
marginiventris and M. autographae parasitized the highest proportion of
hosts in Bermudagrass and paragrass, respectively. FAW larvae parasitized
by A. marginiventris had the smallest mean and range for head capsule
widths indicating that this parasite species permitted the least amount of
host development prior to the host's destruction. Ophion sp. attacked larvae
with the largest h ,ad capsule widths. The tachinids parasitized hosts having
the greatest range of head capsule widths which suggests that these flies
are less influenced by host size and age compared to the hymenopterous para-
sites.
Parasitization levels and FAW abundance were highest in corn during
the months of June and July (Table 2). The first 4 FAW instars in both
corn and Bermudagrass were the most heavily parasitized with substantial
reductions in parasitization occurring in the 5th and 6th instars. The de-
crease in head capsule widths for parasitized and nonparasitized larvae in
corn from April through September may indicate a gradual shift in the
age distribution of the larval populations. The mean head capsule width
for parasitized larvae was always smaller than for nonparasitized larvae.
This condition probably resulted because the hymenopterous parasites,
except Ophion sp., normally destroy FAW larvae in either the 4th or 5th
instar.
The principal parasite on corn was C. insularis, and A. marginiventris
was the principal parasite on Bermudagrass. The differences between
parasites on these 2 hosts may reflect a host plant preference or perhaps a
change in abundance due to time since collections from Bermudagrass were
only made during the last 2 months of the corn collecting period. FAW
larval abundance and parasitization rates were highest in corn during the
months of May, June, and July. The least amount of host development ap-
peared to occur in larvae parasitized by A. marginiventris since these larvae
had the smallest head capsule widths prior to their destruction.
REFERENCES CITED
ASHLEY, T. R., E. R. MITCHELL, N. C. LEPPLA, AND E. E. GRISSEL. 1980.
Parasites attacking fall armyworm larvae, Spodoptera frugiperda, in
late planted field corn. Florida Ent. 63: 136-42.
---- V. H. WADDILL, E. R. MITCHELL, AND J. RYE. 1982. Impact of native
parasites on the fall armyworm, Spodoptera frugiperda, in South
Florida and the release of the exotic parasite, Eiphosoma vitticole.
Environ. Ent. 11: 833-7.
BARFIELD, C. S., J. L. STIMAC, AND M. A. KELLER. 1980. State-of-the-art for
predicting damaging infestations of fall armyworm. Florida Ent. 63:
364-75.
AND J. L. STIMAC. 1981. Understanding the dynamics of poly-
phagous, highly mobile insects. Pages 43-6 in T. Kommedahl, ed. Proc.
Int. Cong. Plant Protection, Washington, DC. Vol. 1. Burgess Publish-
ing Co.
MITCHELL, E. R. 1979. Migration by Spodoptera exigua and S. frugiperda,
North American style. Pages 386-93 in R. L. Rabb and G. E. Kennedy,
eds. Movement of Highly Mobile Insects. North Carolina State Uni-
versity.
Florida Entomologist 66 (2)
MATING BEHAVIOR OF MALACOSOMA DISSTRIA
AT TWO LEVELS OF MATE COMPETITION
DONALD N. BIEMAN AND J. A. WITTER
Insect Attractants, Behavior and Basic Biology Research Laboratory
P. O. Box 14565, Gainesville, Florida 32604 USA
School of Natural Resources, The University of Michigan,
Ann Arbor, Michigan 48109 USA, respectively
ABSTRACT
During a 3-year field study 2 levels of mate competition were observed in
Malacosoma disstria Hiibner, the forest tent caterpillar. Levels were indi-
cated by the mean number per year of male attempts to mate females which
were already in copula (interference rate). Sex ratio at emergence and popu-
lation density probably determined interference rates. In the low level of
mate competition, matings started late in the day and were short in dura-
tion, with some males terminating matings. In contrast, at the high level,
matings began earlier and ended later with some females rejecting males and
some males prolonging matings. The greater mate competition reflected a
higher operational sex ratio, which probably increased the likelihood of
some males finding females earlier and enhanced opportunities for females
to be selective. Furthermore, the chance of sperm competition would be
greater and prolonged matings were likely responses to this. Male terminated
matings probably were responses to low chance of sperm competition.
RESUME
I)urante un studio de campo de 3 afios se observaron 2 niveles de
competencia de aparear en Malacosoma disstria Hiibner. Los niveles fueron
indicados por el promedio por afio de tentativas por los machos de aparear
con hembras ya en c6pula (rango de interferencia). El rango de inter-
ferencia probablemente fue determinado por la raz6n de sexo y por la densi-
dad de la poblaci6n. Al nivel bajo de competencia de aparear, los apare-
amientos empezaron tarde en al dia y duraron corto tiempo, con unos machos
terminando los apareamientos. Al contrast, al nivel alto, los apareamientos
empezaron mas temprano y terminaron mis tarde, con unas hembras re-
chazando los machos, y unos machos prolongando los apareamientos. La
competencia mas intense reflejo una raz6n operacional de sexo mas alta, la
cual probablemente aument6 la probabilidad de que unos machos encontraron
las hembras mds temprano, y aument6 las oportunidades para las hembras
ser selectivas. Ademas, la probabilidad de competencia de esperma seria
mayor, y apareamientos prolongados fueron respuestas probables a esta
condici6n. Los apareamientos terminados por los machos probablemente
fueron resquestas a una probabilidad baja de competencia de espermas.
We examined the mating behavior of' Malacosoma disstria Hiibner, the
forest tent caterpillar (Lepidoptera: Lasiocampidae), in the field at 2 levels
of mate competition. Mate competition was defined as the contest among
males for access to females. The relationship between mating behavior and
intensity of mate competition has been explored in several insect orders: Dip-
tera (Borgia 1980), Hemiptera (McLain 1981, Sill6n-Tullberg 1981), and
Odonata (Pajunen 1966, U6da 1979). Parker (1974) and Wade and
Arnold (1980) have provided theoretical discussions on the subject. This
272
June, 1983
Bieman & Witter: Mating in Malacosoma 273
was the first study to link the intensity of mate competition with mating
behavior in Lepidoptera.
Bieman (1980) described mate search in M. disstria. Males began to
fly from cocoon to cocoon of conspecifics in late afternoon; this behavior
continued into twilight and ceased after dark. Males found females on or
near their cocoons which were wrapped in live leaves of herbs and woody
plants. Although females can emit pheromone (Struble 1970, Percy and
Weatherston 1971, Chisholm et al. 1980), Bieman (1980) observed that
males engaged in close range search did not seem to rely on pheromone for
orientation. Males hovered near any cocoon even though most contained
parasitic flies. Once a female was located, the male hovered at her side. He
then moved back bending his abdomen towards her and pushed its tip in
the direction of her bursa copulatrix. Copulation frequently followed.
Females did not move more than a few cm until twilight when they
flew. The first preparations for female flight occurred when they began
fanning their wings. Wing fanning took place during twilight whether or
not females were mating or had mated. Wing fanning which occurred
during mating was usually the first sign of its end. Females flew away soon
after parting from their mates. Oviposition was concluded within 2 hours
of wing fanning (Bieman 1980). The moths are nonfeeding (Stehr and
Cook 1968). Hodson (1941) discovered that the male and female moths lived
about 5 days. The females usually laid their single egg mass within 3 or 4
hours of emergence but males appeared to be reproductively active through-
out their lives.
Males sometimes attempt to mate with females which are already in
copula (Bieman and Witter 1982). This occurs up until females begin
wing fanning. Late arriving males are called interferers, and the mean
number of times pairs encounter late mating attempts (interference rate)
gives an indication of the intensity of mate competition in a given year. We
compared mating behavior in one year with a high interference rate with
that of 2 years with low interference rates.
METHODS
Mating behavior was observed in 1976, 1977, and 1978. We chose plots in
open areas with small trees and high moth population densities (monitored
by pupal population densities). The 1976 and 1977 plots, approximately
0.25 ha each, were located in an outbreak in Ontonagnon County, Michigan.
The 1976 plot was not reused in 1977 because a plot was found with a higher
population density 4 km to the north. The 1978 plot was also about 0.25 ha
and was located in an outbreak area in St. Louis County, Minnesota. Trembl-
ing aspen, Populus tremuloides Michaux, less than 10 m tall was the pre-
dominant tree species and bracken fern, Pteridium aquilenum (L.), covered
the ground in all plots. Moths were observed during the period of adult
emergence on 6 and 8 July 1976, 22 to 28 June 1977, and 8 to 14 July 1978.
Observation began before male flight, which was the onset of mating
activity, and was concluded at midnight. We scanned plots for solitary fe-
males and mating pairs; one complete scan took about 15 minutes. Once a
pair or solitary female was found it was watched continuously until the
female flew away or midnight, whichever came first. Because of the way we
scanned, some matings went unobserved for up to 15 minutes. The start was
274 Florida Entomologist 66 (2) June, 1983
observed in 6 of 16 matings in 1976, 6 of 10 in 1977 and 3 of 5 in 1978. In the
other cases, the times when the matings were found were used as starts.
We recorded the time of each mating start (start), female wing fanning
(fanning), and mating end (end). The time elapsed between fanning and
end was referred to as prolongation. Its value could be negative; this would
occur if a mating ended before fanning. In 1976, 4 of 16 pairs had not com-
pleted mating at midnight. However, all females had fanned by then. There-
fore in 1976, the means for end and prolongation were underestimated.
We noted: (1) whether or not males held on to the substrate (leaves,
cocoon surfaces, or twigs) during copulation, (2) movement of mating
moths, (3) mating attempts by interferers (also called encounters with
interferers or instances of interference), (4) mate rejections, and (5) the
sex of moth which terminated a mating. We defined one mating attempt by
an interferer as all of the abdominal probes made by the individual until he
departed. A mate rejection referred to any time a male flew away without
mating with a solitary female after she walked away or shielded her bursa
copulatrix from his probing abdomen. The moth which pulled away from or
shook loose from its mate at the end of a mating was considered the mating
terminator.
Time was recorded in CDT and then standardized as min after sunset
for each sample member. Standard deviations are given along with means.
RESULTS
INTERFERENCE RATES
The 1977 and 1978 interference rates, which were not statistically
different, were 3.5 and 10.5%, respectively, of the 1976 value (Table 1).
This indicates 2 different levels of mate competition: high in 1976 and low in
1977 and 1978. Since no differences in any of the mating variables between
1977 and 1978 were found, we combined results for these years (Table 1).
MATING BEHAVIOR IN THE LOW INTERFERENCE YEARS
Matings started late in the day (Table 1) and no females rejected mating
attempts (n = 9). Males held on to the substrate throughout copulations.
Matings ended early; prolongations never exceeded 8 min (Table 1). In fact
in 1977, 2 males terminated matings and flew away. They had mated 34 and
85 min before separating. Their mates remained in the position they main-
tained during mating, respectively, for 117 and 57 min before wing fanning.
This accounts for the negative means for prolongation in 1977 and the
combined totals of 1977 and 1978. The means without the 2 negative values
were 4.1 3.4 min in 1977 and 4.3 3.0 min for the combined total of 1977
and 1978. No other males terminated matings. Interestingly, the male which
ended his mating 57 min prior to fanning had endured interference twice.
He was the only male to encounter interferers in 1977.
MATING BEHAVIOR IN THE HIGH INTERFERENCE YEAR
Matings began early in the day (Table 1) and 4 of 6 females, which
were observed before they mated, rejected mating attempts. These females
moved a few cm away from approaching males or pushed their abdomens to
275
Bieman & Witter: Mating in Malacosoma
00
00 E00' 0
+1 +1 +1 +1 +1
co o(o I
cd 6)
C1 1F00 I I
+1 +1 +1 +1 +1
0 0
oa Ci)
00 00r 00
00 e ';.
E-4
o > .
S '.C 0 + +
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Florida Entomologist 66 (2)
the substrate making their bursa copulatrices inaccessible. Seven mating
males did not hold on to the substrate for the entire time prior to fanning.
We referred to them as hangers and the 9 males which held on the substrate
the entire time before fanning were called holders. The hangers held on by
their claspers an average of 74.0 60.5 min. One female, which previously
rejected a mating attempt, flipped off her mate, which had hung for the
whole 90 min mating, and remated 5 min later. The new mate held on to a
leaf for its entire 93 min mating. Only 1 difference in the mating variables
was found between hangers and holders: the interference rate for hangers
(10.44.5) was greater than that for holders (2.01.9) (Mann Whitney
U-Test, P < 0.05). Eighty-one percent of the interference (8.4 encounters
per mating) occurred while the hangers were suspended. This was 4.2
times as much interference as holders encountered during entire matings.
Thus, it appeared that hanging males were more likely to attract interferers.
Males frequently resisted female efforts to end matings. Seven of 16 pro-
longations were longer than the maximum of 8 min observed during the
low interference years; these ranged from 20 to 140 min. There was no
relationship between the number of interference per mating and length of
prolongation for 1976 (Spearman's rank correlation coefficient = -0.034,
P > 0.1). Three males involved in the long prolongations hung during them.
One of these flew in tandem with the female until landing in another tree
where the mating continued. Four other males also involved in long pro-
longations walked during them in the direction that their mates pulled;
distances walked ranged from 5 cm to more than 50 cm. Walking was ob-
served only once in the low interference years. Considering all years, hang-
ing and walking occurred during prolongation in all 7 matings prolonged
more than 8 min and in only 2 of 24 prolonged 8 min or less. These be-
haviors appeared to reduce the ability of the female to escape the male.
DISCUSSION
Sex ratio and population density probably influence the intensity of
mate competition (Parker 1974, Wade and Arnold 1980). If the sex ratio of
moths increases, the interference rate should also increase. Increases in
population may result in shifts in behavior which would add to the number
of interferers, or would decrease the distance between individuals making
it more likely for searching males to encounter mating pairs. Demographic
conditions were consistent with the levels of mate competition observed in
this study. In 1976, a sex ratio at emergence of 1.65 was determined for
an area within 1 km of our plot (Lorimer, unpublished). In contrast, a sex
ratio of 1.01 was found for the 1978 plot (Bieman 1980). The sex ratio was
unknown for the 1977 plot. The moth population density was at least 10
times greater in 1976 than 1977 and 1978 (Bieman 1980).
The high interference rate in 1976 probably reflected a larger ratio of
searching males to receptive females (operational sex ratio, Emlen and
Oring 1977) than the low rates in 1977 and 1978. A higher operational sex
ratio would increase the likelihood of some males finding females earlier and
the greater number of early starts in the high interference year supports
this.
A higher operational sex ratio would also increase the opportunities for
females to be selective. Mate rejections, which were observed only in the
276
June, 1983
Bieman & Witter: Mating in Malacosoma 277
interference year, support this contention. It was unlikely that rejections
resulted only because females were unreceptive due to previous matings
since females mated within minutes of rejecting males. The basis for the
rejections is unknown (see Rutkowski 1982 for a discussion on mate choice
in Lepidoptera). Hangers provide further evidence for greater female choice
in the high interference year. We believe these males hung to counteract
their mates' efforts to reject them. This reaction, like the similar responses
made by males during prolongation, would reduce the ability of the female
to escape the male. It is probably harder for a female to pull apart from a
male that is not connected to an unmovable object than one that is. This
hanging response did not always succeed. One female did fling off her
hanging mate, but 90 min elapsed before the male fell off.
However, if hanging reduces the chance of rejection, why do most males
hold on to the substrate? We speculate there are costs to hanging. Hanging
may be more energetically costly than holding, and pairs with hangers
attract more interference (probably resulting because of the enhanced con-
spicuousness of these mating pairs), which increases the chances of inter-
ferers inflicting wounds. These wounds can impede or stop reproductive ac-
tivity (Bieman and Witter 1982). Therefore, males may hold on to the sub-
strate unless they are attempting to prolong the period of copulation.
In the lesser wax moth, Achroia grisella (Fabricius) (Lepidoptera:
Pyralidae), males sometimes prolong second matings in order to pass
through a refractory period (Greenfield and Coffelt 1983). This is an un-
likely explanation for the long prolongations observed in 1976 since the
highly competitive situation found in 1976 would not have promoted oppor-
tunities for remating.
Sperm competition, the competition of sperm from 2 or more males for
the fertilization of eggs within a female (Parker 1970), was a likely com-
ponent of M. disstria mating behavior. We propose that long prolongations
occurring during the high level of mate competition were responses to high
probabilities of sperm competition as Sill6n-Tullberg (1981) suggested for
Lygaeus equestris L. (Hemiptera: Lygaeidae). We offer 2 hypotheses of
how prolonged matings would lower the chances of sperm competition: (1)
the longer a male extends a copulation the fewer males his mate encounters
after the mating, (2) the percentage of eggs fertilized increases the longer a
male delays his mate from remating; thus, we expect sperm precedence to
be like that of the eastern spruce budworm, Choristoneura fumiferana
(Clemens). In C. fumiferana, there appears to be a positive relationship be-
tween a first male's success at fertilizing eggs and time passed before a
second male copulates (Retnakaran 1974).
Male terminated matings observed in the low levels of mate competition
may be responses to low probabilities of sperm competition. Males can
fertilize more than 1 female in a day (Stehr and Cook 1968). Therefore, if
the chance of a female encountering a second male is low, males which
terminate their matings may have the opportunity to search for additional
mates in the same evening.
Instead of prolonging matings longer than 8 min or terminating them,
most males responded in an intermediate manner. These males may also be
counteracting the possibility of sperm competition albeit to a lesser degree
than males which prolonged matings past 8 min. Matings prior to fanning
averaged longer than 75 min in all years and over 150 min in 1976. It was
278 Florida Entomologist 66(2) June, 1983
unlikely that all this time, particularly the additional 75 min in 1976, was
used for passing sperm. Some mating time was probably spent guarding
females from remating.
How males determine whether to end matings before fanning, allow fe-
males to break loose soon after, or resist female efforts to terminate matings
is largely unanswered. Males apparently did not decide to prolong matings
on the basis of contact with interferers. There was no relationship between
the number of encounters with interferers and length of prolongations
within the high interference year. Furthermore, the only male to experience
interference in 1977 was one of 2 males which terminated matings. The role
that age, physical condition, and mating experience played in deciding
termination strategy remains unknown.
ACKNOWLEDGEMENTS
We thank K. F. Keyes, C. Keil, and L. Eavy for their help in collecting
data and N. Lorimer for sharing unpublished data. We also thank E. L.
Hover, M. D. Greenfield, T. Burk, C. O. Calkins, J. Sivinski, C. W. Schaefer,
and R. Thornhill for reviews which helped in shaping the final draft of the
manuscript; and L. N. Bundy, L. A. Waisanen, G. W. Fowler, B. S. Low,
F. Slansky, and G. G. Williams who reviewed earlier versions. Work leading
to this publication was funded in part by USDA McIntire-Stennis funds.
REFERENCES CITED
BIEMAN, D. N. 1980. An evolutionary study of mating of Malacosoma
americanum (Fabricius) and Malacosoma disstria Hiibner (Lepi-
doptera: Lasiocampidae). M.S. Thesis. Univ. Michigan 49 p.
--- AND J. A. WITTER. 1982. Mating wounds in Malacosona: An insight
into bedbug mating behavior. Florida Ent. 65: 377-8.
BORGIA, G. 1980. Sexual competition in Scatophaga stercoraria: Size- and
density-related changes in male ability to capture females. Behaviour
75: 185-206.
CHISHOLM, M. D., E. W. UNDERHILL, W. F. STECK, K. N. SLESSOR, AND G. G.
GRANT. 1980. (Z)-5, (E)-7-dodecadienal and (Z)-5, (E)-7 dode-
cadien-1-ol, sex pheromone components of the forest tent caterpillar,
Malacosoma disstria. Environ. Ent. 9: 278-82.
EMLEN, S. T., AND L. W. ORING. 1977. Ecology, sexual selection and the
evolution of mating systems. Science 197: 215-23.
GREENFIELD, M. D., AND J. A. COFFELT. 1983. Reproductive behaviour of the
lesser way moth, Achroia grisella (Pyralidae: Galleriinae);
Signalling, pair formation, male interactions and mate guarding. Be-
haviour (in press).
HODSON, A. C. 1941. An ecological study of the forest tent caterpillar
Malacosoma disstria Hiibner. Minnesota Agric. Exp. Sta. Tech. Bull.
148. 55 p.
MCLAIN, D. K. 1981. Sperm precedence and prolonged copulation in the
southern green stinkbug, Nezara viridula. J. Georgia Ent. Soc. 16:
70-7.
PAJUNEN, V. I. 1966. The influence of population density on territorial be-
havior of Leucorrhinia rubicunda L. (Odon., Libellulidae). Ann. Zool.
Fenn. 3: 40-52.
PARKER, G. A. 1970. Sperm competition and its evolutionary consequences
in the insects. Biol. Rev. 45: 525-68.
Bieman & Witter: Mating in Malacosoma 279
S1974. Courtship persistence and female guarding as male invest-
ment strategies. Behaviour 48: 157-84.
PERCY, J. E., AND J. WEATHERSTON. 1971. Studies of physiologically active
arthropod secretions. IX. Morphology and histology of pheromone-
producing glands of some female Lepidoptera. Canadian Ent. 103:
1733-9.
RETNAKARAN, A. 1974. The mechanism of sperm precedence in the spruce
budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae).
Canadian Ent. 106: 1189-93.
RUTKOWSKI, R. L. 1982. Mate choice and Lepidopteran behavior. Florida
Ent. 65: 72-82.
SILLEN-TULLBERG, B. 1981. Prolonged copulation: A male's 'post-copulatory'
strategy in a promiscuous species, Lygaeus equestris (Heteroptera:
Lygaeidae). Behav. Ecol. Sociobiol. 9: 283-99.
STEHR, F. W., AND E. F. COOK. 1968. A revision of the genus Malacosoma
Hiibner in North America (Lepidoptera: Lasiocampidae) systematics,
biology, immatures, and parasites. United States Nat. Mus. Bull. 876.
321 p.
STRUBLE, D. L. 1970. A sex pheromone in the forest tent caterpillar. J.
Econ. Ent. 63: 295.
UEDA, T. 1979. Plasticity of reproductive behaviour in a dragonfly, Sympe-
trum parvulum Barteneff, with reference to the social relationship
of males and the density of territories. Res. Popul. Ecol. 21: 135-52.
WADE, M. J., AND S. J. ARNOLD. 1980. The intensity of sexual selection in
relation to male sexual behavior, female choice, and sperm precedence.
Anim. Behav. 28: 446-61.
280 Florida Entomologist 66 (2) June, 1983
SCIENTIFIC NOTE
LATE SEASON SOYBEAN LOOPER (LEPIDOPTERA: NOCTUIDAE)
POPULATION REDUCTION BY WESTERN PALM WARBLERS-On 8
October 1973, many small birds were observed in a one-hectare soybean field
which was being sampled weekly to monitor insect pest populations at the
Auburn University Plant Breeding Unit at Tallahassee, AL. The bird
population was estimated to be between 100 and 125, and individuals were
observed to be actively moving through the soybeans in a series of rapid,
short-distance flights, pecking frequently at leaves and stems. When dis-
turbed, they would retreat to nearby trees, but would return within 5
minutes to resume feeding.
Two of these birds were collected on 9 October and returned to the
laboratory for identification and examination of stomach contents. They
were identified by Dr. Julian Dusi, ornithologist in the Auburn Univ. Dept.
of Zoology-Entomology, as Western Palm Warblers, Dendroica palmarum
palmarum (Gmelin) (Passeriformes: Parulidae). This species is known to
migrate northward through Alabama in the spring during March and
April, cross the lower Appalachians, and move up the Mississippi River
Valley into Canada, where its breeding range extends from central Ontario
to British Columbia. This route is reversed from early September to late
October, with the birds eventually reaching Florida, Cuba, the West Indies,
and the Yucatan where they overwinter (A. C. Bent. 1953. Life Histories
of North American Wood Warblers. Smithsonian Institution, USNM Bul.
203, 734 p.). Feeding occurs along the migratory route wherever food
sources are found. Insects constitute the majority of the diet. During migra-
tion, this species has been noted to congregate in areas of an abundant food
supply and to effectively eliminate such food supplies before moving on
(F. M. Chapman. 1970. The Warblers of North America. Appleton, New
York, 306 p.).
In this study, gut analysis revealed large numbers of medium sized soy-
bean loopers, Pseudoplusia includes (Walker), in one stomach and an as-
sortment of loopers and other arthropods, including a chrysomelid beetle, a
lacewing larva, and spiders, in the second. At the time of collection, soy-
bean looper populations, as determined by standard shake cloth sampling,
were recorded at ca. 6.6 larvae/row meter of soybeans, with larval size
classes being distributed from second through fifth instars. Within 3 days,
the larval population was reduced to ca. 0.3 larva/row meter in the sample
field, and no birds remained in the area.
Western Palm Warblers have been recorded previously as taking "cotton
worms" and "celery leaf-tyers" in Florida cotton and celery crops, re-
spectively (A. H. Howell. 1932. Florida Bird Life. Florida Dept. of Game
and Fresh Water Fish, 579 p.). The observations reported here suggest that
they can serve as effective predators of soybean pests as well, under the
proper circumstances. However, no other comparable incident has been ob-
served by the author since 1973. The fortuitous nature of this encounter
suggests that bird predation may, in some circumstances, account for
otherwise unexplainable reductions in insect population numbers. The
relatively short feeding period of several days could easily be missed when
weekly population samples are taken.
In the situation described, impact of predation on soybean looper dam-
Scientific Note 281
age was minimal since pod fill was already complete, and the plants were
in the early leaf senescence stage. In addition, since the soybean looper
does not overwinter in central Alabama, population reduction at this late
date would have little impact on succeeding generations, assuming that the
resident pest population perishes rather than migrates southward. In more
southerly locations where soybeans are produced, late season bird predation
could possibly provide economic benefit. At any rate, utilization of this food
source provided an energy source which aided in the southern migration
of the warblers. Alabama Agricultural Experiment Station Journal
Series #15-82341.-JAMES D. HARPER, Dept. of Zoology-Entomology, Au-
burn University, AL 36849 USA.
Florida Entomologist 66 (2)
BOOK NOTICE
ANNUAL REVIEW OF GENETICS, 1980, Vol. 14, 485 p.
Contents: The Shifting Balance Theory and Macroevolution, SEWALL
WRIGHT; Double Minutes and Homogeneously Staining Regions: Gene
Amplification in Mammalian Cells, JOHN K. COWELL; Genetic Approaches
to the Analysis of Microbial Development, DAVID BOTSTEIN AND RUSSELL
MAURER; Developmental Genetics of Maize, JOHN D. SCANDALIOS; Attenua-
tion in Amino Acid Biosynthetic Operons, CHARLES YANOFSKY AND ROBERTO
KOLTER; Genetic Control of Nitrogen Assimilation in Bacteria, BORIS
MAGASANIK; DNA Uptake in Haemophilus Transformation, SOL H.
GOODCAL; Abnormal Sexual Differentiation in Humans, JOE LEIGH SIMPSON;
Somatic Cell Genetics and Its Application to Medicine, THEODORE T. PUCK
AND FA-TEN KAO; Interaction Between Host and Viral Genomes in Mouse
Mammary Tumors, P. BENTVELZEN; Genetic Defects in Human Purine and
Pyrimidine Metabolism, J. EDWIN SEEGMILLER AND GERRY R. BOss; Low
Level Radiation and Genetic Risk Estimation in Man, CARTER DENNISTON;
T-DNA of the Agrobacterium TI and RI Plasmids, MICHAEL WEBSTER
BEVAN AND MARY-DELL CHILTON; The Genetics of Development in Dictyo-
stelium Discoideum, SUSAN S. GODFREY AND MAURICE SUSSMAN; Strand
Transfer in Homologous Genetic Recombination, CHARLES M. RADDING;
Molecular Genetics of Yeast Mating Type, KIM A. NASMYTH; ROBERT
EDGAR; Indexes-Author; Subject; Cumulative.-JEL
HALFFTER, G., AND W. D. EDMONDS. To be published in 1983. The
nesting behavior of dung beetles (Scarabaeinae), an ecological and evolu-
tive approach. ub. No. 10. Institute de Ecologia, De luxe edition. 184 p. Price:
USA and Canada, $40 US (surface mail included); Mexico $1800 Mexican
pesos (surface mail included); all other countries, $45 US (air mail in-
cluded). 20% discount valid for any order received before 1 September 1983.
Make checks payable to: Instituto de Ecologia. Address all orders and in-
quiries to: Patricia Reidl M.; Instituto de Ecologia; Ap. Postal 18-845;
Deleg. Miguel Hidalgo; 11800 Mexico, D. F.
CONTENTS: Preface. Chapter 1.-The Scarabaeinae. Chapter 2.-The Eco-
logical Evolution of Scarabaeinae. Chapter 3.-Patterns of Nesting Be-
havior in Scarabaeinae: An Overview. Chapter 4.-Evolution of Nesting
Behavior and Sexual Cooperation. Chapter 5.-Nest Construction and
Architecture in Burrowing Scarabaeinae. Chapter 6.-Other Sexual Re-
lationships in Scarabaeinae. Chapter 7.-The Ovary and Nesting Behavior.
Appendix I-Outline/Classification of the Subfamily Scarabaeinae. Post-
script. Appendix II.-Nidification Behavior of Old World Oniticellini
(Coleoptera: Scarabaeidae). Yves Cambefort. Appendix III.-Nesting
Strategies of Three Species of Coprophagous Scarabaeinae in the Sahel
Region of Niger. Daniel Rougon and Christiane Rougon. Appendix IV.-
Commentaries on Recent Literature. Bibliography. Index I.-Subject Index.
Index II.-Taxonomic Index.-CAS
282
June, 1983
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