The

FLORIDA ENTOMOLOGIST

Volume 61, No. 3 September, 1978

CONTENTS

RICHMAN, D. B. AND W. H. WHITCOMB-Comparatiue Life Cycles of Four Species
OF Predatory Stink Bugs .. ... .. .. 113
FAIRCHILD, G. B.-New and Little Known Florida Tabanidae .................. 121
UEBEL, E. C., M. SCHWARZ, P. E. SONNET, R. W. MILLER, AND R. E. MENZER- 4
Evaluation of the Mating Stimulant Pheromones of Fannia canicularis,
F. pusio, and F. femoralis as Attractants ....... ............. .. . 139
MARTIN, P. B., R. L. RIDGWAY, AND C. E. SHUETZE--Physical and Biological Eval-
uations of an Encapsulated Diet for Rearing Chrysopa carnea............ ...... 145
PICKENS, L. G., AND R. W. MILLER-Using Frozen Host Pupae to Increase the
Efficiency of a Parasite Release Program .. ........... ...... 153
PORTER, C. C.-Ecology and Taxonomy of Lower Rio Grande Valley Zethus .... 159
PORTER, C. C.-Ecological Notes on Lower Rio Grande Valley Sphecini ..... 169
WILKINSON, R. C., A. P. BHATKAR, W. J. KLOFT, W. H. WHITCOMB AND E. S. KLOFT
-Formica integra. 2. Feeding, Trophallaxis, and Inter-specific Confrontation
B behavior .... ... .. ... . . . ... .. . .. ... 179
MITCHELL, E. R., E. W. HAMILTON, AND S. MASUDA-A 12-Volt DC Power Pack
for Electrocutor Grid Traps ... .......... 189
GAGNt, R. J.-A New Species of Cecidomyia (Diptera: Cecidomyiidae) Injurious to
Cones of Slash Pine in Florida ... . . .......... 193
Scientific Notes
NOBLE, W. E., ET AL.-Surface Sterilization of Bulb Mites for Phytopatho-
gen Transmission Studies ....... ...... 120
MERKHOFER, R. M.-Firefly Flicker Fusion Frequency . .......... 167
RANDALL, J. B.-An Apparatus for the Observation of Living Immature and
Small Adult Spiders ... ............. ..... 192
Book Reviews .. ... ... .... ..... ... .. 144, 168, 188
Sustaining Associates. ..... .. ...... .......... 195
New Members ........... 196

Published by The Florida Entomological Society

~8LINDEI

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OFFICERS FOR 1977-78

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THE FLORIDA ENTOMOLOGIST is issued quarterly-March, June,
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This issue mailed October 31, 1978

The Florida Entomologist

COMPARATIVE LIFE CYCLES
OF FOUR SPECIES OF PREDATORY STINK BUGS12,3

DAVID B. RICHMAN, AND WILLARD H. WHITCOMB

Department of Entomology and Nematology
University of Florida, Gainesville, Fla. 32611

ABSTRACT
Four species of asopine stink bugs, Alcaeorrhynchus grandis (Dallas),
Euthyrhynchus floridanus (L.), Podisus maculiventris (Say), and Stiretrus
anchorago (Fab.), were reared in the laboratory under identical conditions.
Two groups of each species were maintained; one under simulated Florida
field conditions at a variable temperature ranging from 180-300C, averaging
approximately 260C, and the second at a constant temperature of 27C.
The photoperiod was 14 L:10 D for all groups, and all were fed at the same
time with the same larval instars of the cabbage looper, Trichoplusia ni
(Hiibner), and the soybean looper, Pseudoplusia includes (Walker). At the
given temperatures P. maculiventris and S. anchorago completed their life
cycles in 1 month, while A. grandis and E. floridanus took 2 months. The
egg stages lasted approximately 5 days for P. maculiventris, 6 days for S.
anchorago, 16 days for A. grandis, and 19 days for E. floridanus. All species
were reared through at least 1 generation from eggs laid by laboratory-
reared individuals.

Predatory stink bugs are beneficial insects because they attack a number
of plant feeding lepidopterous and coleopterous larvae (Forbush and
Fernald 1896, Morrill 1906, Whitmarsh 1916, Knight 1923, Plummer and
Landis 1932, Howard and Landis 1936, Hayslip et al. 1953, LeRoux 1964,
Whitcomb and Bell 1964, Oetting and Yonke 1971, 1975, Warren and Wallis
1971, Whitcomb 1973, Waddill and Shepard 1974, 1975, Ables 1975). No
laboratory comparisons of predatory stink bug life cycles have been made
for 2 or more different species reared under the same experimental condi-
tions. Such a comparison should reveal the reproductive strategies em-
ployed by each species. The life cycles of 4 common asopine stink bugs
found in Florida were compared under identical laboratory conditions.
All of the species studied, Alcaeorrhynchus grandis (Dallas), Euthyrhyn-
chus floridanus (L.), Podisus maculiventris (Say), and Stiretrus anchorago
(Fab.), have been reported from soybeans (Whitcomb 1973, Waddill and
Shepard 1974, 1975) and may reduce numbers of lepidopterous pest insects.

MATERIALS AND METHODS
Specimens of A. grandis and S. anchorago were collected in a soybean

This publication was supported in part by the National Science Foundation and the En-
vironmental Protection Agency, through a grant (NSFGB-34718 later known as BMS 75-04223),
to the University of California. The findings, opinions, and recommendations expressed herein
are those of the authors and not necessarily of the University of California, NSF, or EPA.
2 Fla. Agricultural Experiment Station Journal Series No. 1104.
'Hemiptera: Pentatomidae.

Vol. 61, No. 3, 1978

The Florida Entomologist

field near Gainesville, Florida, in October 1977. All field collected
specimens were kept at a variable laboratory temperature ranging from
18'-300C, averaging approximately 260C, with a photoperiod of 14 L:10 D.
The A. grandis nymphs collected in the field were allowed to mature,
mate, and lay eggs in quart jars. Adults of S. anchorage were also kept in
quart jars and allowed to mate and lay eggs. Eggs of E. floridanus and
P. maculiventris were obtained from laboratory cultures where the former
had been maintained through 1 and the latter through many generations.
Eggs of all 4 species were placed in petri dishes in the laboratory or in an
environator (constant temperature of 27 1C.) The bottom of each petri
dish was lined with filter paper and fresh green beans were added 2 to 3
times a week to maintain high humidity. When the nymphs became 2nd
instars they were placed into individual petri dishes and fed caterpillars
of the cabbage looper, Trichoplusia ni (Hiibner), and the soybean looper,
Pseudoplusia includes (Walker), 3 times a week. The nymphs were raised
to adults, and 10 specimens of each nymphal instar were used for measure-
ments of head, anterior pronotum, and humeral width. The time spent in
each nymphal instar was recorded and statistically analyzed. Several
additional egg masses were allowed to hatch and their hatching times re-
corded to obtain more data for comparison.

RESULTS
Development times for the 4 species are presented in Table 1. The egg
stages showed the greatest differences; approximately 16 days for A. grandis,
19 days for E. floridanus, 5 days for P. maculiventris, and 6-7 days for S.
anchorage. The stink bugs differed in the arrangement and numbers of eggs
laid. A. grandis egg masses consisted of 100-200 eggs in multiple rows.
On soybean stems these were in 4-5 rows, but on paper toweling the mass
tended to be broader, up to 10 rows wide. E. floridanus females laid 20-90
eggs in a loose oval mass. P. maculiventris deposited 17-70 eggs, also in a
loose mass. S. anchorago laid 10-23 eggs in a double row. Total develop-
ment time for these stink bugs, from egg to adult, ranged from approxi-
mately 30 days for P. maculiventris and S. anchorago to about 60 days for
A. grandis and E. floridanus. Measurements of the various instars and
adults of the 4 species are presented in Table 2.
Observations of pre-mating behavior indicate that in at least 3 species,
A. grandis, E. floridanus, and S. anchorago, the male mounts over the head
or rear of the female and then maneuvers so that he can take up the charac-
teristic end-to-end copulation position of Hemiptera. The male of S. ancho-
rago buzzes his wings every few seconds during mounting.
All 4 species were observed with natural prey in the field. An A. grandis
4th instar was collected in soybeans feeding on a last instar Anavitrinella
pampinaria (Guen6e) (Geometridae). A 4th instar P. maculiventris was ob-
served consuming a Phoebis sp. larva. An S. anchorago adult was seen
eating a larva of Eurema nicippe (Cramer) (Pieridae), both on Cassia ob-
tusifolia L. in a soybean field. A 5th instar S. anchorago was collected
on goldenrod, Solidago sp., feeding on a lepidopterous larva (undeter-
mined), and an E. floridanus adult was captured eating an adult plant-
feeding stink bug, Euschistus servis (Say), on corn S. anchorago has pre-
viously been reported to eat larvae of the alfalfa weevil, Hypera postica

Vol. 61, No. 3, 1978

Richman and Whitcomb: Predatory Stink Bugs

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TABLE 2. BODY MEASUREMENTS OF IMMATURE AND ADULT SPECIMENS OF
4 SPECIES OF ASOPINE STINK BUGS. (N = 10 IN ALL CASES EX-
CEPT ADULTS).

Species Measurements in mm

Head, incl. eyes

Width SD

Instar

Anterior
pronotum
Width SD

Humeral region

Width SD

Alcaeorrhynchus grandis
1st 0.73
2nd 1.00
3rd 1.48
4th 2.00
5th 2.70
Adult female 3.42
(n = 5)
Adult male 3.01
(n = 5)
Euthyrhynchus floridanus
1st 0.74
2nd 0.90
3rd 1.23
4th 1.66
5th* 2.14
Adult female 2.43

(n = 5)
Adult male*
(n = 5)

2.25 0.03 2.10 0.07 6.40

Podisus maculiventris
1st 0.63
2nd 0.88
3rd 1.25
4th 1.73
5th 2.20
Adult female 2.45
(n = 5)
Adult male 2.33
(n = 5)
Stiretrus anchorage
1st 0.74
2nd 0.91
3rd 1.23
4th 1.60
5th 2.05
Adult female 2.34
(n = 5)
Adult male 2.19
(n = 5)

0.74
0.98
1.40
1.98
2.54
2.63

0.04
0.04
0.06
0.08
0.08
0.08

0.03 2.45 0.05 7.01

0.79
1.03
1.35
'1.81
2.29
2.46

0.00
0.03
0.05
0.08
0.09
0.06

0.03 2.30 0.02 5.58

*Raised in environator at a constant 27"C, 14 L:10 D.

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0.07
0.10
0.12
0.03

0.93
1.26
2.26
3.75
6.45
11.56

0.74
1.06
1.67
2.31
3.17
3.50

0.08 2.97

0.76
0.97
1.28
1.77
2.25
2.28

0.07 9.34

0.01
0.04
0.03
0.05
0.05
0.07

Richman and Whitcomb: Predatory Stink Bugs

(Gyllenhal) (Richman 1977), and Mexican bean beetle, Epilachna varives-
tis Mulsant (Howard and Landis 1936, Waddill and Shepard 1974, 1975).

DISCUSSION
The 4 species of predatory stink bugs studied differed in length of life
cycle, rate of growth, and somewhat in seasonal abundance. Alcaeorrhyn-
chus grandis was found in large numbers in late summer and fall, during
outbreaks of velvetbean caterpillar, Anticarsia gemmatalis Hiibner; E.
floridanus has been collected the year around but is most abundant during
spring and fall in Florida (Mead 1976). Stiretrus anchorage numbers also
peak during spring and fall. Podisus maculiventris was collected through-
out the spring, summer, and fall.
Podisus maculiventris and E. floridanus appear to have the most general
feeding habits (Mukerji and LeRoux 1965, Mead 1976), but all 4 species take
a wide variety of insects. The current study demonstrated that each will
mature and produce viable eggs when fed on 2 species of lepidopterous
larvae. Over 60% of eggs laid by laboratory reared adults of S. anchor-
ago hatched, as compared to less than 25% in the study of Waddill and
Shepard (1974) who fed their specimens exclusively with Mexican bean
beetle larvae. The key element may be provisioning the stink bugs with
more than 1 kind of larva.
In the following discussion our laboratory data are in parentheses after
the data cited. The life history of E. floridanus has been discussed by Ables
(1975) and Getting and Yonke (1975). Ables found that eggs of E. floridanus
took 18.1 (19.1) days to hatch and the nymphal stages lasted 39.5 (39.1) days
at 26.70C and 14:10 photoperiod. Getting and Yonke found that E. floridanus
eggs took 33.4 days to hatch and the nymphal stages lasted 66.9 days at 240C
and 13 L:ll D photoperiod. The cooler rearing temperature and shorter day
length probably account for the long egg and nymphal stages.
Several workers have reared P. maculiventris in the laboratory. The life
history was recorded first by Forbush and Fernald (1896) who reared their
specimens in an insectary in Massachussetts under as close to natural con-
ditions as possible. The eggs took from 7-9 (5) days to hatch and nymphal
development time was at least 25 (22) days. Mukerji and LeRoux (1965)
found that the eggs of P. maculiventris hatched in 4-7 days and that nymphal
development lasted 25-31 days at 270C. Warren and Wallis (1971) found
that the eggs hatched in 6-8 days, and nymphal development took 28.7
days at 210C. Given the temperature difference, our data seem consistent
with published results.
The life cycle of S. anchorage has been discussed by Waddill and
Shepard (1974). At 26.70C with a photoperiod of 14:10, eggs of S. anchorage
hatched in 6.3 (6.8) days and the nymphal stages lasted 18.4 (25.7) days.
The reason for the difference found in, our study is unknown; however, it
is probably not a significant difference.
No studies have been published on the life cycle of A. grandis. We found
that the egg stage lasted 16 days, and maturity was reached 43 days after
hatching.
Podisus maculiventris and E. floridanus were the easiest to rear because
they are prolific egg layers and do well in the laboratory. Podisus tends
to be cannibalistic, but its life cycle is short, and thus as many as 9-10

The Florida Entomologist

generations may be raised in a year. Females may start producing eggs within
1 week of maturing (Mukerji and LeRoux 1965). Euthyrhynchus has limited
cannibalistic tendencies, and nymphs cooperate in prey capture (see also
Ables 1975). Only 4-5 generations could be reared per year because of the
length of its life cycle and because adult females will not lay eggs for
at least 2 weeks after maturing (Ables 1975).
Alcaeorrhynchus grandis nymphs also were observed to cooperate in
prey capture. This species was not as easily reared as E. floridanus, al-
though females produced larger numbers of eggs (100-200) per mass. Egg
masses were not produced as often as those of E. floridanus, and the pop-
ulation level fluctuated more because of mortality in the early instars.
This could be due to genetic load; periodic introduction of field collected
specimens might help maintain the hardiness of the culture. Alcaeorchyn-
chus grandis females reared in the laboratory also took 2 weeks after ma-
turing to produce their first egg masses.
Stiretrus anchorage was the least promising for laboratory rearing. Al-
though it goes through its life cycle very rapidly, females produce small
egg masses (16.4 eggs on the average during this study and 7.1 eggs per mass in
that of Waddill and Shepard 1974) and they start oviposition after 11 days
at 26.7C (Waddill and Shepard 1974).
Further research is needed on the life cycle of these predatory stink bugs
before their role in the control of lepidopterous pests can be fully under-
stood. While life tables have been prepared for some species (e.g. Mukerji
and LeRoux 1969), none has been prepared for A. grandis. More field col-
lected prey records combined with field cage tests would be useful for
evaluating the effectiveness of these species under natural conditions.

ACKNOWLEDGEMENTS
We would like to express appreciation to Drs. Frank Mead, Robert
Crocker, Barbara Saffer, and Reece Sailer for their suggestions, Dr. Robert
C. Hemenway, Jr., for his help with some of the rearing, Dr. D. H. Habeck
for identification of lepidopterous prey, and G. B. Edwards for some prey
records.

LITERATURE CITED
ABLES, J. R. 1975. Notes on the biology of the predacious pentatomid
Euthyrhynchus floridanus (L.). J. Ga. Ent. Soc. 10:353-6.
FORBUSH, E. H., AND C. H. FERNALD. 1896. The gypsy moth, Porthetria
dispar L. Mass. State Printers, Boston. 495 p.
HAYSLIP, N. C., W. G. GENUNG, E. G. KELSHEIMER AND J. W. WILSON.
1953. Insects attacking cabbage and other crucifers in Florida. Bull.
Fla. Agr. Exp. Sta. 534, 57 p.
HOWARD, N. F., AND B. J. LANDIS. 1936. Parasites and predators on the
Mexican bean beetle in the U.S. USDA Circ. 418, 12 p.
KNIGHT, H. H. 1923. Studies on the life history and biology of Perillus
bioculatus Fabricius, including observations on the nature of the
color pattern (Heteroptera, Pentatomidae). 19th Report State Ent.
Minn.: 50-96.
LERoux, E. J. 1964. The application of ecological principles to orchard
entomology in Canada. Can. Ent. 96:348-56.
MEAD, F. W. 1976. A predatory stink bug, Euthyrhynchus floridanus (Lin-

Vol. 61, No. 3, 1978

Richman and Whitcomb: Predatory Stink Bugs

neaus) (Hemiptera: Pentatomidae). Fla. Division Plant Ind. Ent.
Circ. 174, 2 p.
MORRILL, A. W. 1906. Some observations on the spined soldier bug (Podisus
maculiventris Say). U.S. Div. Ent. Bull. (NS) 60:155-61.
MUKERJI, M. K., AND E. J. LEROUX. 1965. Laboratory rearing of a Quebec
strain of the pentatomid predator, Podisus maculiventris (Say)
(Hemiptera: Pentatomidae). Phytoprotection 46:40-60.
MUKERJI, M. K., AND E. J. LEROUx. 1969. A quantitative study of food
consumption and growth of Podisus maculiventris (Hemiptera:
Pentatomidae). Can. Ent. 101:387-403.
OETTING, R. D., AND T. R. YONKE. 1971. Immature stages and biology of
Podisus placidus and Stiretrus fimbriatus (Hemiptera: Pentatomi-
dae). Can. Ent. 103:1505-16.
OETTING, R. D., AND T. R. YONKE. 1975. Immature stages and notes on
the biology of Euthyrhynchus floridanus (L.) (Hemiptera: Penta-
tomidae). Ann. Ent. Soc. Amer. 68:659-62.
PLUMMER, C. C., AND B. J. LANDIS. 1932. Records of some insects predaceous
on Epilachna corrupt Muls. in Mexico. Ann. Ent. Soc. Amer. 25:695-
708.
RICHMAN, D. B. 1977. Predation on the alfalfa weevil, Hypera postica
(Gyllenhal), by Stiretrus anchorage (F.) (Hemiptera: Pentatomi-
dae). Fla. Ent. 60:192.
WADDILL, V., AND M. SHEPARD. 1974. Biology of a predaceous stink bug,
Stiretrus anchorago (Hemiptera: Pentatomidae). Fla. Ent. 57:249-53.
WADDILL, V., AND M. SHEPARD. 1975. A comparison of predation by the pen-
tatomids, Podisus maculiventris (Say) and Stiretrus anchorago (F.)
on the Mexican bean beetle, Epilachna varivestis Mulsant. Ann.
Ent. Soc. Amer. 68:1023-7.
WARREN, L. O., AND G. WALLIS. 1971. Biology of the spined soldier bug,
Podisus maculiventris (Hemiptera: Pentatomidae). J. Ga. Ent. Soc.
6:109-16.
WHITCOMB, W. H. 1973. Natural populations of entomophagous arthropods
and their effect on the agroecosystem. Proc. Miss. Symp. Bio. Control.
Univ. Press of Miss.:150-69.
WHITCOMB, W. H., AND K. BELL. 1964. Predaceous insects, spiders and mites
of Arkansas cotton fields. Ark. Agric. Exp. Sta. Bull. 690, 83 p.
WHITMARSH, R. D. 1916. Life-history notes on Apateticus cynicus and
maculiventris. J. Econ. Ent. 9:51-3.

The Florida Entomologist

SURFACE STERILIZATION OF BULB MITES FOR PHYTO-
PATHOGEN TRANSMISSION STUDIES-(Note). The presence of
breeding populations of Anoetus feronarium Dufour and Rhizoglyphus
robini (Claparede) in association with a complex of bacterial and fungal
pathogens inhabiting the underground storage organs of gladiolus grown in
the sandy soils of Florida suggested possible vector relationships. The in-
teractions and species involved were studied by Engelhard (1969, Phyto-
pathology 59:1025), Poe (1971, Fla. Ent. 54:127-33), and Noble and Poe
(1973, Proc. Fla. State Hort. Soc. 85:401-4).
To determine if mites ingested and excreted phytopathogens, and the
duration of gut infectivity, a technique was needed to destroy or remove
inoculum from the external body surface without harm to the mite or to its
internal flora. Individual mites were removed from lawns of 2 isolates,
Br-ISR (Streptomycin-resistant) and F-1 (an isolate which formed a white
precipitate (WP) on nutrient agar (NA)) of Pseudomonas marginata Mc-
cullough and dipped in 0.78% sodium hypochlorite or 95% ethanol. Mites
remained in the solutions for periods of 3-5 min., were air dried, and placed
on NA for 5 min. Each mite was then aseptically transferred to a nutrient
broth (NB) tube and placed on a rotary shaker at room temperature (760-
78 F). Each NB tube was observed daily for turbidity indicative of bacterial
growth. Samples of turbid broth were streaked on NA and NA + 200 ppm
Streptomycin + 100 ppm penicillin (NASP). Data taken included numbers
of days for turbity to appear in the NB and presence or absence of a white
precipitate, a sensitive test on NA for the F-1 isolate.
No detrimental effects to the mites appeared to result from their treat-
ment with bactericidial solutions or culturing. Observation revealed that
2-3 days were required for turbidity to appear in the NB containing dipped
mites compared to undipped mites, which resulted in turbidity within 24h.
Bacterial colonies with a white precipitate were obtained from the NB cul-
tures containing dipped Rhizoglyphus, indicating that the F-1 isolate of P.
marginata survived in the gut of the mite. Bacterial colonies growing on
the NASP indicated that the streptomycin resistant isolate had also sur-
vived the treatment. Half of the Rhizoglyphus mites tested proved to be
positive for transmission; however, neither isolate of the bacterium was re-
covered from Anoetus. The latter species is a bacterial feeder while the for-
mer is primarily a fungus feeder.
Based on these results, we feel that the surface sterilization techniques
employed in these tests were successful and will be used in our transmission
studies.-W. E. Noble, (Dept. of Ornamental Horticulture, Cal. Poly., San
Luis Obispo); S. L. Poe, (Dept. Entomology & Nematology), and R. E.
Stall (Plant Pathol., University of Florida, Gainesville, 32611).

Vol. 61, No. 3, 1978

The Florida Entomologist

NEW AND LITTLE KNOWN FLORIDA TABANIDAE'2

G. B. FAIRCHILD'

Department of Entomology and Nematology
University of Florida, Gainesville, Fla. 32611

ABSTRACT
Chrysops ifasi n. sp. is described and figured, and males of Chrysops
dorsopunctus Fairchild, C. dixianus Pechuman, C. abatus Philip, C. bistel-
latus Daecke, and C. fulvistigma Hine are described for the first time. Notes
on status and new distribution for these and an additional 12 species of
Chrysops are given. C. upsilon Philip is a new state record. Anacimas gero-
pogon Philip is shown to be a synonym of A. limbellatus Enderlein and the
species is figured. Whitneyomyia beatifica var. atricorpus Philip is confirmed
as a color form. Stenotabanus daedalus Stone is redescribed and figured
from fresh material; the species is placed in Tabanus. Tabanus cayensis
Fairchild is shown to be close to T. campecheanu Townsend from Mexico
and both are figured. New distribution records for 1 species of Hybomitra
and 3 of Tabanus are given.

The notes presented here pertain mostly to specimens and information
accumulated since 1970, partly by the use of a flight trap designed by Dr.
H. V. Weems, Jr., and used by him mostly at Torreya State Park, Liberty
County, and by me at Austin Cary Memorial Forest, northeast of Gaines-
ville, Alachua Co. Other material was taken by hand netting or supplied by
numerous colleagues, who have sent specimens to the Florida State Col-
lection of Arthropods for identification.
I have had the benefit of consultation with Drs. L. L. Pechuman and R. H.
Roberts, most of whose suggestions have been incorporated in the text. The
former also supplied many additional records. I also acknowledge with
gratitude the help of Jayson Glick, who serviced the flight traps at Austin
Cary Forest during my too frequent absences from Gainesville. Finally,
Dr. C. B. Philip has loaned critical material with his usual generosity.
Chrysops abatus Philip 1941, Proc. Ent. Soc. Washington 43:120, female.
Since our notes on this species (Philip et al. 1973) considerable additional
material has turned up, including a single male. This specimen differs con-
spicuously from 2 males of dorsovittatus Hine in lacking thoracic stripes,
in having a broader and less clear cut mid abdominal black stripe and with
duller and brown-tinted lateral pale areas. The fore coxae are black
(yellow in dorsovittatus) as in the females. The apex of the hyaline tri-
angle is broader and blunter in abatus than in dorsovittatus, a condition
also found in females. The wing further differs in having only the apex of
2nd basal cell hyaline, but with an extensive hyaline streak in dorsovitta-
tus. Otherwise the 2 species are very similar. Plesiotype male, Wakulla Co.,

SDiptera: Tabanidae.
2 Florida Agricultural Experiment Station Journal Series No. 943.
Research Associate, Florida State Collection of Arthropods (F.S.C.A.), Gainesville.

Vol. 61, No. 3, 1978

The Florida Entomologist

Florida, Ochlochonee River State Park, 9-IV-77, flight trap, G. B. Fairchild,
coll. Females were quite abundant, taken in a flight trap and netted at this
locality and near McIntyre, across the Ochlochonee River in Franklin
Co. An unusually pale specimen has the dark abdominal markings reduced
to small geminate spots on tergites 3-5, and to a broadly dumbbell shaped
spot on tergite 2, all well separated from each other. This specimen also
has the legs largely yellow, including front coxae, but has the wing pat-
tern of abatus. It is from St. Johns Co., Faver-Dykes State Park, 15-IV-73,
Debra Weems coll.

Chrysops bistellatus Daecke 1905, Ent. News 16:249. The male appears
to be undescribed, though Jones and Anthony (1964) mentioned seeing 1
from Mississippi. As they noted (p. 16), it is easily associated with the female
on wing pattern. The eyes are holoptic, greatly enlarged, so that head is
wider than thorax, the enlarged upper facets clearly differentiated and
demarcated from the small facets and occupying about 2/3 of eye area. The
upper point of the frontal triangle is shiny, and there is a short, median
spur of pollinosity on the frontoclypeus. Color of body and appendages
as in the female, except that 2nd basal cell (M) is 2/3 infuscated. Plesio-
type male, Austin Cary Forest, Alachua Co., Florida, 22-24-V-1975, flight
trap, G. B. Fairchild, coll. The species has been recorded from Florida only
once previously, from Lloyd and Monticello, in Jefferson Co. (Fairchild
1937). It was common at these 2 localities, but taken only for about a
month, 18-IV-21-V. Pechuman (in litt.) also found it abundant at Lloyd
1-2-V-1965. In 1975 it was quite abundant in Austin Cary Memorial Forest
near Gainesville, where 148 females and 1 male were taken in a Weems
flight trap between 28-IV-28-V. Concurrent collections with a hand net
within 500 yards of the trap for about an hour on 14 days between 5-30-V
yielded 432 specimens. D. A. Hurd took specimens 10.5 miles N. of Holt,
Santa Rosa Co., 12-VI-1974 (Pechuman in litt.). In addition, there are 6
females from Blackwater River State Park, Santa Rosa Co., and 1 female
from Camp Blanding, Clay Co., 3-V-1968 in F.S.C.A. The remarkably
short flight season of this species may account for the scarcity of records.
In 1976, in contrast, only 8 specimens were taken in the same trap at the same
location in Austin Cary Memorial Forest, from 10-19-V.
Chrysops brimleyi Hine 1904, Can. Ent. 36:55. Jones and Anthony (1964)
recorded the species only from Jefferson Co., based on Fairchild (1937). Re-
cent collecting showed the species to be common in the northern part of
the state, with specimens at hand from Escambia, Santa Rosa, Okaloosa,
Jackson, Liberty, Wakulla, Franklin, Calhoun, Baker, Alachua, and
Duval counties. It is an early spring species, most records being in March or
April, with a few in May. One of the Duval Co. specimens was taken by D. W.
Anthony, 19-IV-1954, perhaps misdetermined as fuliginosus Wiedemann
and so recorded in 1964.

Chrysops callidus Osten Sacken 1875, Mem. Boston Soc. Nat. Hist.
2:379. Although common and widely distributed in N. America, this species
is uncommon in Florida. Jones and Anthony (1964) saw no material,
merely listing Fairchild's previous records and speculating that McIntyre
might be a misprint for McIntosh in Marion Co. Actually McIntyre is a
small settlement in the Apalachicola National Forest, in Franklin Co.,

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Fairchild: Florida Tabanidae

and I have seen recent specimens from there collected 3-IV-1976 by L. L.
Pechuman. There are also specimens from Escambia, Santa Rosa, Gulf,
Liberty, Leon, and Wakulla counties taken in April and June of various
years. The specimens from St. Augustine and Volusia Co. on which my
records of 1937 were based have been re-examined in M.C.Z. The Volusia
female is dated 11-V-1899, the St. Augustine male undated. There is also a
female labelled Drayton Isl., Florida, 10-V-1897, a locality unknown
to me.
Chrysops carbonarius Walker 1848, List Dipt. Ins. British Mus. 1:203. A
long series was taken by H. V. Weems, Jr., at Torreya State Park, 30-IV-5-V-
1973. These are very dark, the clear areas at tips of basal cells considerably
reduced, the hairs at sides of thorax dull golden rather than whitish, and
the 5th posterior cell (CU1) almost entirely infuscated except for the
small, round, clear spot in its base. These show the same tendency to
melanism as do C. cincticornis nigropterus Fairchild and C. amazon hub-
belli Philip, though less than in the former but more than in the latter. A
specimen from Highlands, North Carolina, seems to be intermediate be-
tween these dark Florida specimens and those from Canada, and there seems
to be no necessity to formally recognize a Florida race. Torreya State Park,
Liberty Co., remains the only locality where this species has been taken in
Florida. Pechuman (personal communication) took it there also in 1965
and 1969.
Chrysops cincticornis nigropterus Fairchild 1937, Fla. Ent. 19:59.
Previously known only from the type locality, Lloyd, Jefferson Co., the
sub-species has been taken occasionally in Torreya State Park, Liberty Co.,
by H. V. Weems, Jr., in April and May of various years, and Goodwin (1976)
recently reported a specimen from Santa Rosa County. In 1975, 2 females
were taken in a Weems flight trap in Austin Cary Memorial Forest, Gaines-
ville, 9-10-IV and 21-22-IV. This represents a considerable southeastern
extension of range. The specimens are even darker than those from Torreya,
the clear spots in distal ends of basal cells are reduced to very narrow,
transverse stripes. The black of the anal and axillary areas is but slightly
less intense than the rest of the wing. Tidwell (1973) felt that Philip's
(1947) raising of this form from variety to subspecies status needed re-
examination. Recent examination of Tidwell's intermediate material from
Louisiana supports his view that nigropterus is no more than an extreme
melanistic form. It is, however, geographically correlated, and the name is
useful for the intensely black specimens from the southeastern periphery
of the species' range. Pechuman (in litt.) reported the form from Chatham
Co., Georgia, and we took it jointly in Franklin and Wakulla Counties in
April 1977.
Chrysops cursim Whitney 1879, Can. Ent. 2:36. Since published Florida
records are in part based on misdeterminations, it may be well to list
the specimens from Florida I have seen, as follows: 5 females Putnam Co.,
Long Lake, 20-25-V-1971, light trap, H. Bishop, coll.; 1 female Putnam Co.,
Melrose, 21-V-1971, J. Butler, coll.; 2 females Alachua Co., 3 mi. E. of
Micanopy on State Road 346, 14-VI-1973, Fairchild and Wilkerson, colls.;
2 females Alachua Co., Univ. Florida Hort. Sta., 10-25-V-1973, Wilkerson,
coll. (Fig. 2); 3 females Sumter Co., Panasoffkee, 27-VI-1973, Carl Scheck,
coll.; 1 female Highlands Co., Archbold Biological Station, Lake Placid,
17-V-1961, H. E. and M. A. Evans, colls. (L. L. Pechuman det.); 3 females

The Florida Entomologist

Alachua Co., Austin Cary Memorial Forest, Gainesville, 7, 12, 20-V-1975,
netted, J. Glick, coll.; 2 females same locality, 2-5, 13-14-V-1975, flight
trap, Fairchild and Glick, colls.; 2 females same locality, 28-V-1976, flight
trap, Fairchild and Glick, colls. These records suggest that cursim, though
sometimes taken with ifasi n. sp., has an earlier main flight season and
different ecological preferences. One of the specimens from Long Lake
listed above was compared and found to agree with topotypes of cursim in
M.C.Z. I also have the remains, abdomen and 1 wing, of a specimen from
New Jersey collected by Daecke and determined as cursim by Whitney.
The eye pattern from a female collected at the University of Florida
Horticultural Unit, Gainesville, Fla., 10-V-73, by R. Wilkerson is illus-
trated (Fig. 2).
Chrysops dixianus Pechuman 1974, J. New York Ent. Soc. 82:185-7.
This recently described species was abundant in collections made with a
Weems flight trap in Austin Cary Forest, near Gainesville, Alachua Co.,
388 females and 1 male being taken between 6-V-31-VII-1975, and 158
females taken between 10-V-13-VIII-1976. It appears to have a shorter
flight season than either C. flavidus Wiedemann or C. reicherti Fairchild
which were taken in the same trap from 8- and 25-IV and 8-9-IX, respec-
tively. The single male agrees closely with the females in color and pat-
tern of thorax, abdomen, and wings, and in leg and antennal color. The
eyes are holoptic, the upper enlarged facets well differentiated from the
small, occupying about 2/3 of eye area, but not sharply demarcated from
the small facets, there being a gradual increase in size extending over sev-
eral rows of facets. Compared to a male of C. reicherti, the wing pattern
easily separates the 2, and in addition the separation of the 2 sizes of eye
facets is more abrupt in reicherti. Males of C. flavidus, in addition to dif-
ferent wing pattern, have swollen antennal scapes and pedicels and a pale
pollinose triangle on 2nd tergite. The plesiotype male of dixianus is la-
belled Alachua Co., Gainesville, Austin Cary Forest, flight trap 24-25-V-
1975, G. B. Fairchild.
Chrysops dorsopunctus Fairchild 1937, Fla. Ent. 19:59. This species was
originally proposed as a variety of fulvistigma Hine 1904, based on 27 fe-
males from Lloyd, Jefferson Co., taken from 21-V-23-VI-1935. I also had
seen specimens from Georgia in the U.S.N.M. In addition to the solid black
truncated triangle on the 2nd tergite, the ocelli of fulvistigma are sur-
rounded by a roughly square, black, shiny patch, while in dorsopunctus
this area is usually largely pollinose in unabraded specimens. In dorso-
punctus the mid and fore femora are often darkened at least at base and
apex, while in fulvistigma they are usually clear yellow. Comparisons of
measurements of height and width of frons, height and width of callus,
and indices derived from these measurements do not indicate clearly sig-
nificant differences, though in the small sample available, the frons and
callus of dorsopunctus are generally slightly wider.
A male plesiotype, Bay Co., Econtina Creek, Highway 20, 8-VI-1975,
K. J. Tennessen, coll., is easily associated with the female on color of
abdomen, although the black median band is wider and more parallel sided
than in most females, and the sides of tergite 3 are irregularly yellowish.
The face is protuberant, shiny black with a shallow median groove which is
yellowish gray pollinose and extends from antennal bases to base of pro-
boscis. The eyes are greatly enlarged, head wider than abdomen, the upper

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Fairchild: Florida Tabanidae

enlarged facets clearly demarcated and differentiated from the small
facets and occupying about 2/3 of eye area. This area in the dried specimen
is light tan, with all borders dark brown, and a dark brown band about 4
facets wide extending from just below ocellar tubercle to lower posterior
corner of large facetted area. The wings are almost wholly hyaline, only
faint traces of the weak crossband being visible. Two of the females of dorso-
punctus from Alachua Co. have the median black figure on tergite 2 an-
teriorly dilute and irregularly tridentate, quite different from the 2-
pronged figure of fulvistigma. Their ocelli are not surrounded by a bare
patch, and I do not consider them intermediates between dorsopunctus
and fulvistigma.
Tidwell (1973), with a series of over 100 specimens from several
localities in southeastern Louisiana before him, found about 25% belonged
to the dorsopunctus form, and there were specimens he considered inter-
mediate. Both forms were taken together, and in view of the intermediates,
he felt he was dealing with a single variable species. But the differences
in the males, and the absence of intermediates in Tidwell's material, now
at F.S.C.A., make it seem more likely that we are dealing with a species pair
similar to Chrysops niger Macquart and C. caluus Pechuman and Teskey.
Perhaps the larvae or pupae may throw more light on the problem. In
the meantime it seems better to treat the 2 forms as separate species until
their true relationship can be discovered, rather than ignore the question
by retaining the nomenclaturally invalid varietal standing for dorso-
punctus.
I have seen dorsopunctus from Alachua, Jefferson, Liberty, Bay, Oka-
loosa, and Santa Rosa counties, with dates ranging from 9-V-5-VII.
Chrysops fulvistigma Hine 1904, Can. Ent. 36:55. Originally described
from North Carolina, the species was first reported from Florida, without
more definite locality, by Philip (1950). Fairchild (1937) did not collect
it in Florida, but reported it from Oxford, Mississippi. Brennan (1935) re-
ported the species from Georgia and Louisiana, but had at least 1 specimen
of dorsopunctus Fairchild in his material. The specimens reported from
Louisiana (Keachie and Osborn) by Hine (1907) appear also to belong to
dorsopunctus, judging from his description, and as noted by Fairchild
(1937:59). Jones and Anthony (1964) saw no Florida specimens. Tidwell
(1973), reporting on the Tabanidae of Louisiana, felt that dorsopunctus
could not be separated from fulvistigma due to the occurrence of intermedi-
ates, and reported both forms as occurring together in the southeastern part
of that state. Tidwell's material, now in F.S.C.A., shows 250+ specimens
of fulvistigma and 16 of dorsopunctus. All specimens were easily assignable
to 1 or the other of these taxa.
In Florida the 2 forms seem to behave as separate species, and all ma-
terial I have seen can be separated on several characters, as detailed under
dorsopunctus. Florida specimens of fulvistigma available to me are all
from west of Tallahassee, as follows: Liberty Co., Torreya State Park,
many females 13-15-VI-1974, 1 female 4-VII-1965, 2 female 13-15-VI-1966,
all Weems, coll. 7 females 20-21-VI-1973, Fairchild and Wilkerson, colls.
1 female Liberty Co., near Bristol, 26-VIII-1946; 1 female Eglin A.F.B.,
Okaloosa Co., no date.
Although Brennan (1935) mentioned seeing a male of fulvistigma, he did
not describe it, merely saying it is like the female and easily associated.

The Florida Entomologist

A male plesiotype of fulvistigma from Oxford, Mississippi, VII-1934, no
coll., now in M.C.Z., is much like the male of dorsopunctus in leg color
and structure and proportions of the head and its appendages. It has the same
pollinose midfacial stripe also. It differs most notably in having the wing
pattern quite strongly pigmented, the dark markings on the pale tan, upper
eye facets broader and heavier, and the abdominal pattern quite different.
The 1st abdominal tergite is yellow, the middle 1/3 blackish, the dark por-
tion with diffuse borders. The 2nd tergite has the middle 1/3 or slightly
more black, with a narrow, middorsal, yellow triangle 2/3 the length
of the segment, widest at the anterior border of the segment, tapering pos-
teriorly to a sharp point. Third tergite largely black, with a yellow,
inverted, middorsal triangle nearly length of segment and broader than
the triangle on 2nd tergite; sides of tergite narrowly dull yellowish.
Fourth tergite blackish brown, with a small dull yellow, inverted triangle
less than 1/2 the length of the segment. Remaining tergites blackish.
Chrysops geminatus Wiedemann 1828, Auss. Europ. Zweifl. Ins. 1:205.
Jones and Anthony (1964) saw no Florida specimens of this species, only
listing the record of Fairchild (1937) from Jefferson Co. Since then, the
species has been taken abundantly at Torreya State Park, Liberty Co.
by Howard Weems, Jr., at Bristol, Liberty Co., and in Wakulla Co., in May
and June of various years. M. A. Tidwell also took the species abundantly
near Cantonment, Escambia Co., 26-27-V-1972, while L. L. Pechuman has it
from Okaloosa Co., VI-1974, C. L. Smith coll.
Chrysops hinei Daecke 1907, Ent. News 18:143 and C. beameri Brennan
1935, Bull. Univ. Kansas 36:265. These 2 very similar species can be easily
confused, and Philip (1955) suggested that both may be infra-specific var-
iants of C. sequax Williston 1887. In Florida, 2 forms seem to be present,
differing consistently only in the pattern of the abdomen as follows:

Chrysops beameri
Abdomen with slender, continuous, black, dorso-lateral stripes from
1st to 5th tergites. Paired dorsal stripes nearly always parallel and con-
tinuous.

Chrysops hinei
Abdomen either without dorsolateral black stripes or these represented
by short dashes, rarely present on 1st and 2nd tergites. Paired dorsal stripes
usually of slightly oblique dashes interrupted by the sutures, rarely
completely parallel and continuous.

The wing pattern is nearly identical, though in beameri the hyaline
triangle seems slightly wider on the average. Both species fly late in the
year, August to October, rarely at other times, and have been taken in the
same general areas, though not surely together. Jones and Anthony (1964)
reported only a single collection of beameri from Alachua Co., taken
5-X-1940. One of these is before me, courtesy of Dr. Anthony, and agrees with
my interpretation of beameri based on material compared with a paratype
in M.C.Z. The specimens from Monticello, Florida, reported as hinei by

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Fairchild: Florida Tabanidae

me (Fairchild 1937) are mostly beameri, but not all. I have also seen
2 females from Blackwater River State Park, Santa Rosa Co., 7-VIII-1976,
Fairchild coll., and Santa Rosa Co., 22-VIII-1955 which I think are beameri.
One has a nearly black callus, but is determined beameri by Stone.
C. hinei was present in small numbers at Austin Carey Forest near
Gainesville from 27-VIII-4-XI-1976, both in a flight trap and netted flying
about the collector. I also took a few in Monticello, Jefferson Co. in
IX-1935, while Jones and Anthony (1964) reported the species from Alachua,
Lafayette, Lake, Liberty, and Marion counties. To this may be added a
female from Clay Co., 26-IV-1973, given to me by a student. L. L. Pechuman
wrote that he has specimens from Highlands, Polk and Columbia counties.
In addition there are also specimens in F.S.C.A. from Manatee, Volusia,
Hillsborough, Pasco, and Bay counties so that the species seems to be
widely distributed in Florida. Its comparative rarity in collections appears
to be due to restricted choice of habitats as discussed by Jones and Anthony
(1964:20), coupled with an unusually late flight season when, due to ex-
cessive heat, few entomologists are in the field.
Chrysops ifasi Fairchild, NEW SPECIES
(Fig. 1, la, Ib)
A pale grey and yellow fly without black markings on abdomen and
with narrowed and pale brownish crossband on wing.
Female. Length 7 mm; of wing 6.5 mm. Frons slightly higher than wide;
index 1.26, slightly wider at base than vertex, black in ground color, thinly
grey pollinose. Ocellar tubercle pollinose, except for small bare spots
outward of each ocellus. Frontal callus yellow, not shiny, wider than high,
obtusely pointed in middle of upper and lower margins. Subcallus, fronto-
clypeus, and genae yellow in ground color, the pollinose areas light yel-
lowish, the bare areas shiny. Antennae (Fig. la) with segments, beginning
with scape, having the proportions 15:10:31, so that 3rd segment is longer
than preceding 2 combined. Style slightly longer than basal portion.
First 2 segments dark yellow, shiny, quite inflated, clothed with black
hairs, apical 1/2 of 2nd segment brown. Basal portion of 3rd segment dark
yellow, the style black, contrasting. Palpi and theca of proboscis yellow,
shiny, the former with sparse black hairs, labella black.
Mesonotum steel grey, very faint median and paired sublateral darker
stripes visible in certain lights. Pronotal lobes dull yellow. Notopleural
lobes dark grey. Pleura unstriped, grey, slightly paler than mesonotum.
Scutellum grey at base, yellow at apex, whole thorax pollinose, without
bare areas. Legs, including coxae, yellow, except for dusky apical 1/3 of
fore tibiae and apical 2 or 3 tarsal segments of all legs. Wings as figured
(Fig. 1), the crossband and apical spot more extensive than in Chrysops
delicatulus Osten Sacken and cursim Whit., about as in pudicus Osten
Sacken, but very pale brownish.
Abdomen orange yellow in ground color, paler on 1st 2 or 3 segments,
slightly darker terminally. Each segment from 2 to 5 inclusive, has a pair of
poorly defined, semitransparent, dorsolateral spots which appear slightly
darker. The surface is subshiny, beset with sparse yellowish hairs on hind
and lateral borders of tergites, which form low median triangles on the hind
borders; remainder of tergites sparsely black haired. Ventrally abdomen
wholly dull yellow, sparsely yellow haired and not so shiny as dorsum.

The Florida Entomologist

'. lb-- .' **- -^ - "~~ -

Fig. 1. Chrysops ifasi Fairchild n. sp. Wing, paratype. Monticello, Jef-
ferson Co., Florida 27-IX-35, G. B. Fairchild, coll.; (a) antenna, holotype;
(b) eye patterns of holotype and paratype.
Fig. 2. C. cursim Whitney. Eye pattern, female, University of Florida
Horticultural Unit, Gainesville, Fla., 10-V-73, R. Wilkerson, coll.

Holotype female, Alachua Co., Florida, 3 miles east of Micanopy on
State Road 346, 14-VI-1973, pine flatwoods, Fairchild and Wilkerson,
colls. The species is named for the Institute of Food and Agricultural
Sciences of the University of Florida, which has supported this work.
The eye pattern (Fig. Ib) was drawn for the holotype and 1 paratype.
It is most similar to that of pudicus as figured by Daecke (1906) and of
cursim (Fig. 2), but the dark pattern is considerably more reduced, the pos-
terior margin having but faint traces or none of dark.
Paratypes (all Florida): 1 female, same data as holotype; 15 females,
Monticello, Jefferson Co., 21-VIII-, 11-, 12-, 16-, 18-, 19-, 27-IX-1935, G. B.
Fairchild coll.; 1 female, Greenville, Madison Co., 24-VIII-1935, G. B.
Fairchild coll.; 1 female, Fruitland Park, Sumter Co., V-1921; 1 female,
Withlacoochee State Forest, Sumter Co., 19-VII-1973, Fairchild and Wil-
kerson, colls.; 1 female, Ocala National Forest, Marion Co., 11-VII-1973;
Fairchild and Wilkerson, colls.; 1 female, Ochlochonee River State Park,
Wakulla Co., 22-VI-1973, Fairchild and Wilkerson, colls.; 1 female, 2 mi.
N.W. Orange Springs, Putnam Co., 27-VIII-10-IX-1975, insect flight trap,
J. Wiley, coll.; 1 female, Okaloosa Co., Holt Fish Hatchery, flight trap,
5-7-VIII-1976, G. B. Fairchild, coll.; 8 females, Smith Ranch, Umatilla,
Lake Co., 26-VII-1954, on man (2) 3-IX-1953, on horse (6), Jones and An-
thony, colls. One of the 1953 specimens was det. C. cursim by Stone, sev-
eral others were det. cursim by Anthony, so that this appears to have been,
at least in part, the species recorded as cursim in Jones and Anthony (1964).
The specimens they listed from Dixie and Putnam counties, as well as the
3 reared females are not available to me, so that I cannot state whether
they were this species or cursim. Finally, Dr. L. L. Pechuman lent me a
single female, Floyd's Island, Okefenokee Swamp, Georgia, 17-VIII-1945,
Fattig, coll. This was det. cursim by Stone and recorded as cursim by Fattig
(1946).
The paratypes are somewhat variable, a few showing distinct dark stripes
on mesonotum, and rarely the base of discal cell may be hyaline. The
scutellum may be all grey or all orange yellow. The abdomen may show
distinct traces of pollinose yellow, middorsal triangles, and very rarely
there may be a faint brownish darkening of the dorsolateral, semitrans-
parent spots.

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Fairchild: Florida Tabanidae

Holotype and some paratypes to be deposited in F.S.C.A., other para-
types in M.C.Z., Cornell, U.S.N.M., and collections of L. L. Pechuman,
C. B. Philip, and the author.
This species usually is readily separable from cursim by the lack of, or
fainter, thoracic stripes, absence of black on abdomen, and broader cross-
band and apical spot. From pudicus it differs in lacking any black on the
abdomen, usually lacking thoracic stripes, and having the wing pattern
dilute brownish. Most pudicus also have the frontal callus at least nar-
rowly black bordered above. I suspect that ifasi is most closely related to
pudicus, while cursim is more related to delicatulus. Each may be a diluted
expression of the earlier described species of the pair.
Both cursim and pudicus have been taken together with ifasi, but I have
seen no intergrades. Chrysops delicatulus is recorded as far south as South
Carolina, where it is known only from Cheraw State Park, Chesterfield Co.,
according to Pechuman, who wrote me that this is a disjunct population
and the species is not otherwise known south of Maryland. A specimen
from Gainesville, Alachua Co., Florida, 19-VIII-1929, E. R. Jones, coll.,
was det. delicatulus by C. B. Philip, but it is intermediate between cursim and
delicatulus. The callus is yellow, as are legs, especially fore coxae, and it
seems to me much better placed as a dark variant of cursim.
Chrysops montanus Osten Sacken 1975, Mem. Boston Soc. Nat. Hist.
2:282. This species was reported from Jefferson Co. by Fairchild (1937),
the record repeated by Jones and Anthony (1964), but re-examination of 2
of these specimens in M.C.Z. shows them to belong to the following form.
Chrysops montanus perplexus Philip 1955, Rev. Brasiliera Ent. 3:111-2,
(Fig. 7d). This subspecies was based on 1 male from North Carolina and 1
male from Orange Co., Florida. Subsequently Pechuman (1957) described
the female from Welaka, Putnam Co., Florida. It differs from northern
montanus chiefly in being slightly darker, the hind femora generally
largely black, and the apical spot more extensive, often filling whole wing
apex and leaving the hyaline triangle as a narrow stripe bordering the cross
band. Intermediate specimens are not uncommon, even in Florida, so that
the form is probably the end of a dine, like the dark forms of C. carbon-
arius Walker, amazon Daecke, and cincticornis Walker. It hardly seems to
warrant even a varietal name. In addition to the localities listed by Jones
and Anthony (1964), I have seen specimens from Leon, Jefferson, and Dixie
counties. The species was fairly common in Austin Cary Forest near Gaines-
ville, Alachua Co., in 1975 and 1976, 124 females being taken in traps
and netted between 29-IV-27-V-1975 and 29-III-10-V-1976.
Chrysops nigribimbo Whitney 1879, Can. Ent. 11:35. Though reported
from Florida by Philip (1947), Jones and Anthony (1964) recorded it only
from Alachua Co. Specimens in F.S.C.A. and my collection are from Es-
cambia, Santa Rosa, Wakulla, Alachua, Baker, Columbia, and Clay
counties, with dates from 23-IV (Alachua Co.) to 7-VIII (Escambia Co.).
Chrysops pudicus Osten Sacken 1875, Mem. Boston Soc. Nat. Hist. 2:381.
This variable species has been difficult to determine, and literature reports
are often unreliable. I now have material compared with the lectotype in
M.C.Z., as well as specimens det. by L. L. Pechuman, J. Bequaert, and
C. B. Philip. Based on these specimens, the species can be characterized as
follows. Third antennal segment subequal to or shorter than sum of 1st
2 segments, the latter slender or slightly inflated. Mesonotum and scutel-

The Florida Entomologist

lum dark steel grey, with at least traces of a narrow median and dorso-
lateral darker shiny stripes. Rarely the scutellum is tipped with yellow.
Frons grey pollinose, nearly always clearly higher than wide. Ocelli on
a pollinose tubercle, with only small bare spots or none. Callus usually
yellow in center and below, the upper and lateral margins brown to black,
but occasionally all yellow or all black. Facial callosities entirely
yellow. Dark crossband of wing clear cut, slightly wider than length of
discal cell, reaching hind margin only in 4th posterior cell (3rd M), with
a separate, light infuscation along Cu2. Outer margin nearly straight, apical
spot not or but slightly drop shaped, reaching to wing apex, not obscuring
more than distal 1/2 of R4, usually less. Hind femora brown to black,
mid and fore femora wholly or largely yellow. Abdomen yellow on 1st
2 tergites, the 1st with a small brown to black patch beneath scutellum,
the 2nd with an inverted, broadly U-shaped, black mark whose apex reaches
or nearly reaches anterior margin of the segment, and whose lateral arms
generally fail to reach the hind margin. The size and shape of this figure
vary considerably, being small and clear cut to large and somewhat dif-
fuse, rarely with extensions laterally paralleling the hind border of tergite
2. Third and succeeding tergites largely dark, brown to black, with a pale
hind border of variable width which may form a median triangle, acute
to broad and obtuse; lateral borders wholly pale. Rarely there may be
a little yellow on dorsolateral areas of tergites 3 and 4. Ventrally the
abdomen is largely pale, but with median dark spots increasing in size from
tergites 3 to 6.
The species most likely to be confused with pudicus in Florida is C. dim-
mocki Hine. This is usually a more robust species, the frontal callus nearly
always black, the crossband strongly bowed outwardly, the apical spot
clearly drop shaped, and the median black figure on tergite 2 more inverted
V shaped, very often with lateral posterior extensions along the hind margin
of the tergite.
Specimens are before me from Alachua, Gulf, Broward, Wakulla, Jef-
ferson, Manatee, Dade, Lafayette, Highlands, and Jackson counties, with
dates from 2-IV-1971, Highlands Co., to 27-IX-1935, Jefferson Co.
Chrysops tidwelli Philip and Jones 1962, Fla. Ent. 45:67. This species
has been known hitherto only from the types from Escambia Co., taken
in August 1960. I made a trip in early August 1976 to the same area, but
was unsuccessful in securing specimens. I did, however, collect a single
female at Blackwater State Park, Santa Rosa Co., 7-VIII-1976, netted while
flying about my head along a trail through pine woods.
Chrysops upsilon Philip 1949, Ann. Ent. Soc. America 42:458-60. Al-
though described from Georgia and taken abundantly in Louisiana (Tidwell
1973), this species was not mentioned by Jones and Anthony (1964), nor was
Florida included in its range by Philip (1965), so that the following records
appear to be the 1st for the state: 4 females, Liberty Co., Torreya State
Park, 20-VI-1973, Fairchild and Wilkerson, colls.; 1 female, Liberty Co.,
Apalachicola River near Bristol, 14-VI-1972, P. H. and N. F. Carlson,
colls.; Washington Co., Pine Log Creek at East River, 16-X-1970, P. H.
Carlson, coll. L. L. Pechuman (in litt.) added Jackson Co., 31-V-1965,
F. T. Moore, coll. The specimens recorded by Fairchild (1937) as "uni-
vittatus Macquart" have been re-examined. They are C. macquarti Philip,

Vol. 61, No. 3, 1978

Fairchild: Florida Tabanidae

as surmised by Jones and Anthony (1964:21), rather than the true univit-
tatus Macquart.
Anacimas limbellatus Enderlein 1925, Mitt. Zool. Mus. Berlin 11:376
(Fig. 3, 4). In my opinion the specific name should date from 1925, as the
1923 reference given in the North American catalogue (Philip 1965) is a very
brief definition of the genus, so that limbellatus was a nomen nudum at that
time. I have now been able to study a total of 31 females, 5 males of
this species, as listed here: (1) 1 male, Florida, Andreas Bolter collection,
det. Tabanus gracilis Wiedemann? by C. W. Johnson, in C. B. Philip col-
lection. This specimen was reported as A. geropogon by Philip (1952).
(2) 1 male Springfield, Florida, 21-IV-1951, ex pitcher plant, C. B. Philip,
coll. This specimen is in poor condition, greased, and with wings tattered;
it is the "Allotype" of limbellatus, so labelled and designated by Philip
(1952). (3) 1 male Alachua Co., 2 mi N.W. Gainesville, Florida, 16-II-1975,
in G. B. Fairchild collection. (4) 1 female, Carolina Beach, North Carolina,
20-IV-1932, holotype of A. geropogon Philip in C. B. Philip collection.
(5) 1 female, Jacksonville, Florida, Englehart, coll., general, alcoholic,
in C. B. Philip collection. (6) 1 female, Dog Island, Florida, 17-X-1947,
alcoholic reported as geropogon by C. B. Philip 1952, in C. B. Philip col-
lection. (7) 1 female, Dewees Island, South Carolina, IV-1929, in M.C.Z.
This specimen is closest to type of geropogon. It was det. limbellatus by
Bequaert and bears a note on and sketch of the eye pattern done by him.
The eye sketch is essentially like the eye figured in the present paper. I
did not draw the antenna. (8) 1 female, Hancock Co., Mississippi, 3-IV-1975,
R. H. Roberts, coll., det. limbellatus by Roberts. It lacks antennae, but
except for minor differences in frontal characters and paler wings seemed
the same as Florida specimens. (9) 1 female Florida, Franklin Co., Wright
Lake, Apalachicola Natl. Forest, 21-IV-1971, flight trap, Weems and Fair-
child, colls., wing length 9 mm, the smallest specimen measured. (10)
5 females, Alachua Co., Florida, Austin Cary Memorial Forest, N.W. of
Gainesville, 27-III-7-IV-1975, flight trap, G. B. Fairchild, coll. (11) 17 fe-
males, same locality, 1-III-12-IV-1976, flight trap, G. B. Fairchild, coll.
(12) 1 female, same locality as No. 9, 23-III-1973, Weems, coll., in F.S.C.A.
Some of the above specimens were reported by Goodwin (1976). Drawings of
eye pattern of 1 specimen from No. 11, is given here, as well as of frontal
characters of No. 11, and antennae of Nos. 1, 2, 4, 5, 6, 9, and 10 (all 5).
As can be seen from the figures, there is wide variation in the antennae,
even within the series from Austin Cary Forest, so that little reliance
can be placed on this character to separate geropogon and limbellatus. Fron-
tal characters are also variable, but not correlated and not very striking.
The type of geropogon seemed to be paler, more red sided, and the dark mid-
dorsal abdominal band more prominent than in my fresh Florida speci-
mens. The same was true as to the Dewees Island, South Carolina specimen,
but both are over 40 years old and the color differences noted may be due
to fading. All except the Dog Island, Florida specimen were taken in Feb.,
March or April, and it is 1 of the earliest species on the wing. Dr. L. L. Pechu-
man showed me additional specimens on a recent visit, viz.: 1 female,
Carteret Co., North Carolina, Gales Creek, 10-VI-1971, M. A. Tidwell,
coll.; 1 male, Southern Pines, North Carolina, IV-1907; 1 male, Echolls
Co., Georgia, 11-IV-1970, Beshear, coll.; and 1 female, Wakulla Co., Flor-
ida, Ochlochonee River State Park, 4-IV-1976, in CO, baited trap, L. L.

The Florida Entomologist

, -3

\Ib v'
3'~:~

4 /

4f

7

7

-I
II1 1

t '

49

- .L

4. 4d

49 4

6 1

4I ^S,^4 <

4

49-A

'n
/'7

i)

8

i',1,

Fig. 3. Anacimas limbellatus Enderlein, female. Fla., Austin Cary
Forest, Gainesville, 1-3-III-76; (a) eye pattern, white=reddish green;
stippled=dull green; black=purple; (b) frons, antenna, palpus; (c) an-
tenna at greater magnification.
Fig. 4. Anacimas limbellatus Enderlein. Morphological variation in an-
tennae of some specimens listed in the text. (a) male, Fla., Andreas Bolter
collection; (b) male, Fla., Springfield, 21, IV-1951, ex pitcher plant, C. B.
Philip, coll.; (c) female, N. C., Carolina Beach, 20-IV-1932, holotype of

Vol. 61, No. 3, 1978

Fairchild: Florida Tabanidae

Pechuman, coll. More recently Pechuman, Goodwin and I took 5 males,
33 females at the last locality during the second week in April 1977.
I conclude that there seem to be no firm characters for separating A.
geropogon Phil. 1936 from A. limbellatus End. 1925 (New synonymy). The
species ranges from Mississippi to North Carolina, south to central Florida.
As to the position of Anacimas, it is placed in the Tabanini by Philip
in 1947, but in the Diachlorini in 1965. The specimens before me have the
basicosta unusually reduced in size, a small usually rounded knob, which
may be entirely bare or variably setose, with the setae sometimes as dense
as on adjoining costa. The frons is unusual in the great development of
the median callus, which is more protuberant than in most other North
American species, or in any Neotropical species. The vertex in the female is
slightly inflated, but with a short median bare streak. In the male, there is
a well marked tubercle between the eyes, though without vestiges of ocelli.
The proboscis is unusually short, the labella small and pale, its tip barely
exceeding the palpi. There seems no information on the biting habits, and
the species may not be haematophagous, though Goodwin (1976) felt his
specimen may have been attempting to bite. In any event, the species is
not obviously Diachlorini, and I suspect that its diachlorine characters may
be secondary, though until the larvae can be studied there seems no point
in trying to place it. Anacimas dodgei (Whitney), the only other species in
the genus, has a similar basicosta.
Whitneyomyia beatifica (Whitney) 1914, Can. Ent. 46:353 and var.
atricorpus Philip 1950, Ann. Ent. Soc. America 43:122. Although described
as a subspecies, Philip stated he believed atricorpus was only a melanistic
variety, and reduced it to varietal status in 1965. Tidwell (1973) reported
only the atricorpus form from Louisiana, while Philip (1950) reported 55
of the typical form from Apalachicola, Florida. There are in F.S.C.A. 1
female from the Apalachicola series reported by Philip and 2 females,
St. Theresa, Franklin Co., 17-IV-1938, of the typical form, and 1 female,
Mimmsville, Georgia, 24-V-1907, and 5 females, Umatilla, Lake Co.,
Florida, 4-V-1951, on horse, C. M. Jones, coll., of the dark form. In my col-
lection there are 4 females of the typical form from Austin Cary Forest
near Gainesville, 12-14-V-1975 and 3-V-1976, and the following specimens
of atricorpus: 5 females, Austin Cary Forest, Gainesville, 22-24-IV-1975,
5- and 23-IV- and 12-V-1976; 1 female, Umatilla, Lake Co., 4-V-1951, C. M.

A. geropogon Philip; (d) female, Fla., Jacksonville, Englehart, coll., (e)
female, Fla., Dog Island, 17-X-1947; (f) female, Fla., Franklin Co., Wright
Lake. Apalachicola Natl. Forest. 21-IV-1971, Weems and Fairchild, colls.;
(g) females, Fla., Alachua Co., Austin Cary Memorial Forest, N.W. Gaines-
ville, 27-III-7-IV-1975. G. B. Fairchild, coll.
Fig. 5. Tabanus campecheanus Townsend. Cancun, Quintana Roo,
Mexico, Rosel, coll. Palpus, frons, antenna.
Fig. 6. Tabanus cayensis Fairchild. Florida, Cedar Island, Taylor Co.,
Weems, coll. Palpus, frons, antenna.
Fig. 7. Tabanus daedalus Stone. Florida, Austin Cary Forest, Gaines-
ville. 8-IX-75, J. Glick, coll. Palpus, frons, antenna.
Fig. 8. Tabanus daedalus Stone, holotype, palpus, frons, antenna; (a)
antenna of specimen from Jacksonville, Fla. mentioned in text; (b) eye
pattern of specimen from Austin Cary Forest, Gainesville, Fla., 6-VIIII-76.
White = greenish bronze; black = dull brown.

The Florida Entomologist

Jones, coll.; 1 female Shady Lake, Gainesville, Florida, 11-VI-1953, on
mule, D. W. Anthony, coll.; 1 female, Alachua Co., Hatchett Creek,
11-IV-1974, K. J. Tennesen, coll. There thus does not seem to be any geo-
graphic separation of the 2 forms, and the species appears to be dimorphic
in respect to the abdominal hair colors.
Hybomitra difficilis (Wiedemann) 1828, Auss. Zweifl. Ins. 1:165. This
species was known in Florida only from a single collection by H. V.
Weems, Jr., at Torreya State Park, Liberty Co., 12-IV-1960; it was first
reported by Philip (1961) and repeated by Jones and Anthony (1964). Since
then a long series of both sexes has been taken there by Weems in April
and May in various years and by Pechuman in 1969. Frank Mead took
a female, Gainesville, Alachua Co., 20-III-1964, labelled "edge of ham-
mock," and in 1974, 1975, and 1976, I took a short series in hardwood forest
at the University of Florida Horticultural Station, 23-27-III-1975, 12-22-
III-1976, and 2-IV-1974. Harold Greenbaum took specimens at Florida
Caverns State Park, Jackson Co., 25-26-III-1972. L. L. Pechuman also took
specimens in a CO,-baited canopy trap in Wakulla Co. in April 1977. Hybo-
mitra difficilis thus appears to be an early spring species favoring deciduous
hardwood forest. Greenbaum's and my specimens were taken in flight traps,
as was much of Weems' material.
Stenotabanus daedalus Stone 1938, U.S.D.A. Misc. Publ. 305:32 (Fig. 11).
Although described from Gainesville, Florida, this has remained one of the
most rarely collected and least known of North American Tabanidae.
Stone (1940) reported the 2nd specimen from 8 mi. N.W. of Gainesville.
Philip (1941) described the male (from Volusia Co., Florida, 6 mi. W. of
Barberville, 6-IX-1938) and noted that it was not a Stenotabanus. He sug-
gested that it might be better placed in Glaucops Szilady on account of the
2-segmented style, but the type species of that genus, hirsutus (Villers),
has a bare basicosta and very different frons according to Chvala, Lyneborg,
and Moucha (1972:397), although 3 males and 2 females seen by me have
the basicosta sparsely setose. Philip (1947) placed daedalus in Glaucops, but
in 1965 he returned it to Stenotabanus. P. W. Fattig (1946:14), listed the
species from Fort Valley, Georgia, Aug., and Reidsville, Georgia, Sept.
Jones and Anthony (1964) reported 2 females taken from a horse in Lake
Co., Florida. I have studied the holotype from Gainesville, Florida, 26-IV-
2-X-1914, now in A.M.N.H. and the following additional material: 1 fe-
male, Lee's Ranch, Lake Co., Florida, 17-VIII-1951, horse, A. J. Graham,
coll., no doubt 1 of the specimens reported by Jones and Anthony (1964) as
it was given me by the latter. 1 female, Jacksonville, Florida, 15-IX-1943,
T. Aitken, coll., labelled as comp. with type by L. L. Pechuman Sept. 1960,
and det. 1949 as Glaucops daedalus by C. B. Philip. 1 female, Alachua Co.,
Austin Cary Memorial Forest, 8-IX-1975, in truck, J. Glick, coll. 1 female
same locality, 19-IX-3-X-1975, flight trap, J. Glick, coll. 23 females
same locality, 2-, 6-VIII-, 3-24-IX-, and 8-X-1976, flight trap, G. B. Fair-
child, coll., and CO,-baited Malaise trap, R. Roberts, coll.
As can be seen by the accompanying figures, (Fig. 7, 8) there are no un-
usual characters, other than the reduced number of segments in the 3rd
antennal segment, which would warrant separation of this species from
Tabanus, and it is herewith placed as Tabanus daedalus (Stone) NEW
COMBINATION. Based on the specimens I have seen, the species may be
redescribed as follows: Female, length 9-11.5 mm, of wing 7.5-9 mm. Eyes

Vol. 61, No. 3, 1978

Fairchild: Florida Tabanidae

bare, greenish bronze, with 3 narrow dark bands, as figured, very similar to
eyes of T. zythicolor Philip. Frons nearly parallel sided, index 4.3 to 4.5
dark yellowish-grey pollinose, as is subcallus. Frontal callus dull
yellow, rounded to subquadrate, well separated from eye margins, nar-
rowly or barely attached to the small and irregular, brown to black,
median callus. No tubercle, bare or discolored patch or markedly dif-
ferentiated hairs at vertex, the postocular fringe of short inconspicuous
pale hairs. Antennae dull yellowish brown, the scape and pedicel neither
enlarged nor notably hairy, the style slightly darker than basal plate,
with 2 or 3 annuli. Palpi moderately inflated basally, the apex rather
slender, white with mixed black and white hairs. Frontoclypeus and genae
pale grey, white haired, except upper margins between antennal bases
and eyes, which are brownish and dark haired.
Mesonotum blackish, grey pollinose, sparsely golden haired, the noto-
pleural lobes paler, dark haired. Pleura and coxae dark grey, white haired.
Legs dull yellowish brown, mostly black haired, the apices of fore tibiae
and all tarsi slightly darker, but tibiae not strongly bicolored. Wings
with venation normal, no appendix at fork, greyish hyaline, veins yellow
to brown, the costal cell distinctly yellowish. Basicosta with setae about
as dense as on adjoining costa.
Abdomen mainly dull yellowish brown in ground color, with an ill-
defined, darker, median triangle on tergites 1 and 2, mainly sparsely dark
haired, but with small, paired, dorsolateral spots, hind and lateral mar-
gins of at least tergites 1 to 4, and small median triangles on all tergites,
sparsely yellow haired. Pollinosity accentuates the hair colors, pale under
pale hairs, darker under black hairs, but the pattern is in any case not
very pronounced or clear cut.
All recorded specimens were taken in August, September or October,
and from a relatively small area from central Georgia to north central
Florida, eastward to the coast. The short late flight season and inconspic-
uous appearance of this species may have caused it to be overlooked or
confused with some member of the longus group, or even taken for a dwarf
specimen of pallidescens Philip or fulvulus Wiedemann.
Tabanus cayensis Fairchild 1935, Fla. Ent. 18:53-54 (Fig. 6). This small
species has been seldom collected, and the male remains unknown. Com-
parison with the, until recently, poorly known Tabanus campechianus
Townsend 1897 (Can. Ent. 29:197) from the Yucatan peninsula of Mexico,
shows the 2 to be very similar. The frontal indices of cayensis range from
2.50-2.96, mean of 6 specimens= 2.64, while 4 specimens of campechianus
range from 2.36-2.67, mean=2.47. These differences hardly seem to be
significant. Since campechianus has never been figured, I give here figures of
both species for comparison (Figs. 5, 6). The more slender palpi and broader
antennae of cayensis, together with the dark coxae and femora will prob-
ably serve to separate the species. It seems likely that the 2 species had a
common ancestry not too far in the past, and in fact subspecific status
for cayensis might better reflect their supposed relationship. However,
since males of neither are known, it seems better to defer making any change
until further evidence accrues. I have seen, in addition to the types, the
following specimens of cayensis, all females and all from Florida: 1, Big
Pine Key, Monroe Co., 7-V-1961, Weems, coll., Philip, det.; 1, Collier
Co., 14-V-1949, Fairchild det.; 1, Big Pine Key, 26-V-1950, J. S. Harger, coll.,

The Florida Entomologist

Pechuman det.; 1, Cedar Island, 16 mi. N.W. Steinhatchee, Taylor Co.,
16-V-1969, Weems, coll., Philip det.; 1, Bonita Springs, Lee Co., 15-IV-1975,
Vingst, coll., Fairchild det.; the Everglades, 25-IV-1928, Fattig, coll.,
Pechuman det.; Stock Island, 18-VIII-1975, Pierce, coll., Pechuman det.;
Adams Beach, Taylor Co., 14-V-1965, Pechuman, coll. et det.; Marco Beach,
Collier Co., IV-1955, H. E. Evans, coll., Pechuman det.; Cape Coral, Lee
Co., 8-V-1964, Pechuman, coll. et det.; Sanibel Island, Lee Co., 1-5-VI-
1973, A. Lewis, coll., Pechuman det.; and same loc., 11-V-1973, W. Wirth,
coll., Fairchild det. The species appears to be restricted to the Gulf coast
of Florida and the lower keys, but may be more widely distributed, being
easily confused with dwarf specimens of nigrovittatus Macquart unless
studied carefully. The latter has a quite different and rarely overlapping
frontal index, 2.93-4.17, mean of 29 specimens = 3.44.
I have studied only 5 specimens of campechianus, as follows: 1 fe-
male, Corozal, British Honduras, 15-V-1960, J. Strangways-Dixon, coll.;
1 female, Mexico, Campeche, 48 mi. N. Puerto Real, 22-VI-1966; 2 females,
Cancun, Quintana Roo, Mexico, 24-25-VII-1973, Pletch and Rosel, colls.,
Philip det. (Fig. 5); 1 female, Campeche, Mexico, V-1963, F. S. Blanton,
larger and yellower than the others. The Corozal specimen was relaxed
and shows a green eye with 1 slender, faint, median, dark stripe.
Tabanus nigrovittatus fulvilineis Philip 1957, American Mus. Novit.
1858:3-6, (Fig. 2). This subspecies was based primarily on the color and
pattern of the abdomen, the median as well as dorsolateral stripes said
to be yellow, while in nigrovittatus the median stripe is said to be gray.
This character is difficult to assess in denuded specimens, since the under-
lying pollinosity seems always grey, only the hairs varying in color, and
then not much. I first thought that the 2 forms could be separated on
color of the coxae and femora, as was done by Jones and Anthony (1964:70),
yellow in fulvilineis, black in nigrovittatus, and indeed pale specimens
do look different, with little or no underlying black on abdomen, and ab-
dominal hair stripes broad and yellowish. These color characters, however,
are unsupported by structural differences, the frontal indices and degree
of divergence of the frons in a series of 18 pale-legged specimens, including
2 Florida paratypes, not being significantly different from the same indices
taken from 29 black-legged specimens, including 2 Bahama paratypes,
and specimens from Massachusetts, North Carolina, New York, and Mexico,
as well as Florida. Philip's holotype had a narrower frons than any of the
4 paratypes seen, or than any other specimen measured by me. I give here
the ranges and means of the measurements and indices I took. Pale-legged
specimens: frontal length 43-57, mean=49.83; basal width 12-16, mean=
13.67; vertex width 14-17, mean = 15.22; frontal length/ basal width (frontal
index) 3.21-4.08, mean=3.65; vertex width/basal width (divergence index)
1.00-1.29, mean=1.12. A divergence index of 1.00 indicates a completely
parallel-sided frons. Black-legged specimens: frontal length 38-56,
mean=49.79; basal width, 11-18, mean= 14.48; vertex width, 13-19, mean=
16.03; frontal index 2.93-4.17, mean=3.44; divergence index, .94-1.29, mean
=1.11. Measurements were taken by measuring the image of the frons as
seen through a camera lucida with a millimeter rule. I have seen pale-
legged specimens only from the Gulf coast of Florida, from Pinellas Co.
to Okaloosa Co. Philip (1957) recorded pale-legged specimens from Ship
Island, Mississippi, and Hernando Co., Florida, as well as within the range

Vol. 61, No. 3, 1978

Fairchild: Florida Tabanidae 137

given above. All material from the east coast of Florida, as well as the
Keys and southern tip of the state, has been black-legged, as is the Bimini,
Bahamas material reported by Philip. Unfortunately, too many specimens
are intermediate in regard to leg color and cannot be clearly placed on
this character. In general, the paler, more yellow specimens are smaller
and may more often have yellow or brown frontal calli. I conclude that
fulvilineis probably represents a paler, more yellowish moiety of a more
yellowish southern population of nigrovittatus, which I, at least, cannot
consistently separate. The name remains available should future studies
reveal biological differences indicating reproductive isolation of this form.
Tabanus subsimilis Bellardi 1859, Saggio Ditt. Messicana 1:66. Under
the name of Tabanus vittiger schwardti Philip, Jones and Anthony (1964)
recorded this species as having been taken in Bay, Escambia, and Dade
counties, although they themselves seem not to have seen Florida speci-
mens. One of the paratypes of T. vittiger schwardti was said to be from
Everglades Exp. Sta., Dade Co. (Philip 1942), while in 1952 Philip recorded
males and females taken in pitcher plants (Sarracenia flava Linnaeus) in
the outskirts of Panama City (Bay Co.). I have not traced the source of
the Escambia Co. record in Jones and Anthony. Although I was inclined
to query these records, especially the 1 from Dade Co., feeling that there
might have been confusion with T. vittiger guatemalanus Hine, I have now
seen an undoubted female specimen shown to me by L. L. Pechuman. It
was collected at Pomona Park, Putnam Co., 19-V-1961, by A. and H. Deit-
rich, and is now at Cornell Univ. Both Pechuman (in litt.) and I are still
sceptical of the Dade Co. record, though accept provisionally those from
N.W. Florida.
Tabanus vittiger guatemalanus Hine 1906, Ohio Nat. 7(2):24. As suggested
by Philip (1957:7), this appears to be a coastal form throughout its ex-
tensive range in the Caribbean and Central America as well as in Florida.
Jones and Anthony (1964) listed the species from Dade, Monroe, and
Pinellas counties, and Pechuman (in litt.) recorded specimens from Collier
Co., while a specimen from Naval Air Station, Jacksonville (Duval Co.),
June 1969, taken at porch light by L. W. Teller furnishes a surprising ex-
tension of range. The specimen is in excellent condition and seems un-
likely to have been a stray brought in from further south by aircraft.

LITERATURE CITED
BRENNAN, J. M. 1935. The Pangoniinae of Nearctic America (Tabanidae,
Dipt.). Bull. Univ. Kansas 36(8): Univ. Kansas Sci. Bull. 22(13):
249-401.
CHVALA, M., L. LYNEBORG, AND J. MOUCHA. 1972. The horseflies of Europe.
p. 1-500, + 8 pl. Copenhagen.
DAECKE, E. 1906. On the eye coloration of the genus Chrysops. Ent. News
17:39-42, Pl. 1.
FAIRCHILD, G. B. 1937. A preliminary list of the Tabanidae (Dipt.) of Flor-
ida. Fla. Ent. 19:58-63; 20:10-1.
FATTIG, P. W. 1946. The Tabanidae or horseflies and deerflies of Georgia.
Emory Univ. Mus. Bull. 4:1-26.
GOODWIN, J. T. 1976. Notes on some "rare" eastern Nearctic Tabanidae
(Dipt.); state records and host-parasite relationship for other species.
Fla. Ent. 59:63-6.

The Florida Entomologist

HINE, J. S. 1907. Second report upon the horseflies of Louisiana. Bull.
93, Agric. Exp. Stat. Louisiana State Univ., p. 1-59.
JONES, C. M. AND D. W. ANTHONY. 1964. The Tabanidae (Dipt.) of Florida.
Techn. Bull. 1295, A. R. S., United States Dept. Agric., p. 1-85.
PECHUMAN, L. L. 1957. Descriptions of Tabanidae previously known from
one sex only. Bull. Brooklyn Ent. Soc. 52:29-34.
PHILIP, C. B. 1941. Comments on the supra-specific categories of Nearctic
Tabanidae (Dipt.). Can. Ent. 73:1-14.
PHILIP, C. B. 1942. Notes on Nearctic Tabanidae. Part III. The Tabanus
lineola complex. Psyche 49:25-40.
PHILIP, C. B. 1947. A catalogue of the blood-sucking fly family Tabanidae
(horseflies and deerflies) of the Nearctic region north of Mexico.
American Midl. Nat. 37:257-324.
PHILIP, C. B. 1950. Corrections and addenda to a catalogue of Nearctic
Tabanidae. American Midi. Nat. 43:430-7.
PHILIP, C. B. 1950a. New North American Tabanidae (Dipt.). Part II. Tabani-
dae. Ann. Ent. Soc. America 43:115-22.
PHILIP, C. B. 1952. Notes on tabanid flies and other victims caught by the
carnivorous plant, Sarracenia flava. Fla. Ent. 35:151-5.
PHILIP, C. B. 1955. New North American Tabanidae IX. Notes on and keys
to the genus Chrysops Meigen. Rev. Brasiliera Ent. 3:47-128.
PHILIP, C. B. 1957. New records of Tabanidae in the Antilles. American
Mus. Novitates 1858:1-16.
PHILIP, C. B. 1961. Coop. Econ. Ins. Rept. 11(5):48.
PHILIP, C. B., in Stone et al. 1965. A catalogue of the Diptera of America
north of Mexico. United States Dept. Agric. Handbook No. 276
(Tabanidae, p. 319-42).
PHILIP, C. B., H. V. WEEMS, Jr., AND G. B. FAIRCHILD. 1973. Notes on
eastern Nearctic Haematopota, Merycomyia and Chrysops, and de-
scription of male of C. zinzalus (Diptera:Tabanidae). Fla. Ent.
56:340-6.
STONE, A. 1940. Two new Nearctic Tabanidae and some new records and
corrections (Diptera). Proc. Ent. Soc. Wash. 42:59-63.
TIDWELL, M. A. 1973. The Tabanidae of Louisiana. Tulane Studies in
Zoology and Botany 18(1-2):1-95.

Vol. 61, No. 3, 1978

The Florida Entomologist

EVALUATION OF THE MATING STIMULANT
PHEROMONES OF FANNIA CANICULARIS, F. PUSIO,
AND F. FEMORALIS AS ATTRACTANTS.2,3

E. C. UEBEL4"7, M. SCHWARZ', P. E. SONNET",
R. W. MILLER4, AND R. E. MENZER7

ABSTRACT
The respective mating stimulant pheromones of Fannia canicularis
(L.), F. pusio (Wiedemann), and F. femoralis (Stein) were evaluated for
their effectiveness in increasing the trap catch of a sugar bait. The phero-
mones of F. canicularis [(Z)-9-pentacosene] and of F. pusio [(Z)-ll-hentria-
contene] produced a slight, but significant increase in capture of males
but not of females when compared with an untreated sugar bait. The phero-
mone of F. femoralis [(Z)-11-hentriacontene] did not increase the trap catch
of either sex.

Mating stimulant pheromones produced by female flies have been
identified for the house fly, Musca domestic L. (Carlson et al. 1971,
Rogoff et al. 1973, Uebel et al. 1976); the face fly, Musca autumnalis
De Geer (Uebel et al. 1975b, Sonnet et al. 1975); and the stable fly, Sto-
moxys calcitrans (L.) (Muhammed et al. 1975, Uebel et al. 1975a, Sonnet
et al. 1977).
Recently, some of us (Uebel et al. 1977, 1978a, and 1978b) investigated
the cuticular lipid constituents of Fannia spp. and identified the com-
ponents that were most active in sexually stimulating male F. canicul-
aris, F. pusio, and F. femoralis. The external lipid of F. canicularis females
was found to contain a large amount of (Z)-9-pentacosene, which serves
as the mating stimulant pheromone for the males of this species. Male
F. canicularis were found to contain a large amount of an unusual cutic-
ular component that was identified as 8-heneicosanol acetate (Uebel et
al. 1977). The monoolefin (Z)-11-hentriacontene was identified as the most
active compound in stimulating copulatory responses from males of F.
pusio and F. femoralis (Uebel et al. 1978a, 1978b).

Diptera: Muscidae.
2A portion of a dissertation intended for submission by the first author to the Graduate
School of the University of Maryland in partial fulfillment of the requirements for the Ph.D.
degree.
3This paper reports the results of research only. Mention of a pesticide in this paper does
not constitute a recommendation for use by the U.S. Department of Agriculture nor does it
imply registration under FIFRA as amended. Also, mention of a proprietary product in this
naper does not constitute a recommendation or an eAdorsement of this product by the USDA.
SChemical and Biophysical Control Laboratory, AEQI, FR, SEA, USDA, Beltsville, Md.
20705.
5 Biologically Active Natural Products Laboratory, AEQI, FR, SEA, USDA, Beltsville,
Md. 20705.
6 Present address: Insect Attractants, Behavior, and Basic Biology Research Laboratory, FR,
SEA, USDA, Gainesville, Fla. 32604.
SDepartment of Entomology, University of Maryland, College Park, Md. 20742.

Vol. 61, No. 3, 1978

The Florida Entomologist

Carlson and Beroza (1973) and Morgan et al. (1974) have reported that
the addition of the house fly sex attractant pheromone, (Z)-9-tricosene
(muscalure), to a trap will significantly increase the catch of both male
and female house flies. This paper reports the results of our evaluation
of the mating stimulant pheromone of these 3 species of Fannia as attrac-
tants under simulated field conditions.
MATERIALS AND METHODS
Tests with F. canicularis and F. femoralis were conducted in a 3 x 3 x 3
m-room that contained a battery of 20 White Leghorn hens. Since under
similar conditions so few F. pusio were captured, the tests reported here
were conducted in a screen cage (107 x 81 x 97 cm) in a test room of the
Fly Control Laboratory. Attractancy was determined by comparing the
catch of a trap baited with sugar with that of a trap baited with sugar plus
the test material.
Two traps, constructed from 1.4-liter plastic containers (C-70, Tristate
Plastics, Henderson, KY) with inverted entrance cones, were placed at op-
posite sides of a 71-cm-diam circular platform. They were supported 3 cm
above the platform by 3 metal legs. The platform, which was 25 cm above
the floor of the test cage and 48 cm above the floor of the test room,
was rotated at 2 rpm by a 1/20 hp electric motor (Model 3M125, Dayton
Electric Co.).
The baits were prepared by saturating circles of filter paper (127 mm,
Grade 617, Easton-Dikeman) with 4 ml of petroleum ether containing the
test material or with 4 ml of petroleum ether alone. After the solvent
had evaporated, the filter paper was pressed into a petri dish. Then 1.5 g of
sucrose was sprinkled onto the filter paper and the dishes were placed
beneath the traps. Except that a concentration of 1 mg was omitted from
the tests with F. femoralis, all test compounds were evaluated at concen-
trations of 25, 5, and 1 mg/filter paper. In addition, a 1:1 mixture of (Z)-11-
hentriacontene and tricosane was tested at 10 mg to determine whether this
combination, which stimulates mating of F. femoralis males, would be ef-
fective as an attractant for this species.
Tests were made by releasing ca. 1500 recently closed flies from the
laboratory colony into the test room or cage every other day. The traps
were collected after 24 h, and the captured male and female flies were
counted. Then, the traps were washed (also alternated over treated and un-
treated filter papers), and the petri dishes were replaced with clean ones
containing fresh filter paper, sugar, and test material. The numbers of males
and females captured during 10 or 15 such trapping periods in the traps
with and without test material were individually analyzed by the 2-way
analysis of variance.
RESULTS
Table 1 shows that the trap over' the filter paper treated with (Z)-9-
pentacosene (25, 5, or 1 mg) captured more F. canicularis males than the
trap over the untreated filter paper. Also, these traps captured more F.
canicularis females than the controls though the difference was only
significant for a concentration of 5 mg.
When 8-heneicosanol acetate, a cuticular component of male F. cani-
cularis, was tested at 1 or 5 mg on filter paper, the numbers did not sig-
nificantly differ from those captured in the trap without this material.

Vol. 61, No. 3, 1978

Uebel et al.: Fannia Mating Stimulants

TABLE 1. NUMBERS OF F. canicularis CAPTURED IN 2 TRAPS, 1 BAITED
WITH SUGAR PLUS (Z)-9-PENTACOSENE AND THE OTHER WITH
SUGAR ALONE.*

Mean number captured
Catch
Amount of Trap with Trap with ratio:
(Z)-9-pentacosene treated untreated treated/ Level of
on filter paper filter paper filter paper untreated significance

MALES
1 mg 70.4 40.2 1.8 1%
5 mg 53.6 28.4 1.9 1%
25 mg 53.3 32.1 1.7 1%
FEMALES
1 mg 37.8 25.8 1.5 N.S.
5 mg 27.8 17.7 1.6 5%
25 mg 22.2 18.9 1.2 N.S.

*Means are for ten 24-h collections at each concentration.
Catches of F. pusio males (Table 2) were greater in traps baited with
sugar plus 25, 5, or 1 mg (Z)-ll-hentriacontene than in traps baited with
sugar only. However, (Z)-ll-hentriacontene had no apparent effect on the
catches of F. pusio females.
There were no significant differences between the numbers of F. femoralis
males captured in traps baited with sugar plus 25 or 5 mg of (Z)-ll-hen-
triacontene and in traps baited with sugar alone (Table 3). Also, there
was no significant difference between the catches of F. femoralis females in
traps over the untreated filter paper or filter papers treated with 5 mg. In
the test with 25 mg of (Z)-ll-hentriacontene, the traps over the untreated
paper captured more F. femoralis females, an indication that the females
were repelled by this concentration of the monoolefin.
Table 3 shows that the alkane plus monoolefin combination had little
effect: ca. the same numbers of male and female F. femoralis were captured
in the trap over the sugar and test mixture as over the sugar bait alone.

DISCUSSION
Our present study demonstrates that male F. canicularis and F. pusio
are moderately attracted by their respective mating stimulant pheromones,
while both (Z)-9-pentacosene and (Z)-11-hentriacontene are essentially in-
effective in attracting females. Interestingly, Carlson et al. (1971) also
found that (Z)-9-tricosene attracted maje house flies in laboratory tests,
though in field tests (Carlson and Beroza 1973), nearly equal numbers
of the sexes were attracted.
We feel that the main function of the mating stimulant pheromones
borne by female flies is to initiate sexual or courtship responses from con-
specific males. Thus, the attraction they demonstrate is probably a second-
ary effect. Since these pheromones were effective in only slightly increasing
the trap catch of the males of 2 of the Fannia species, their use as an
attractant for the control of these pest flies remains doubtful.

The Florida Entomologist

TABLE 2. NUMBERS OF F. pusio CAPTURED IN 2 TRAPS, 1
SUGAR PLUS (Z)-11-HENTRIACONTENE AND THE
SUGAR ALONE.*

BAITED WITH
OTHER WITH

Amount of
(Z)-11-hentriacontene
on filter paper

Mean number captured

Trap with Trap with
treated untreated
filter paper filter paper

MALES

1 mg
5 mg
25 mg

1 mg
5 mg
25 mg

60.9
24.9
16.9

52.7
31.1
16.3

FEMALES

*Means are for fifteen 24-h collections at each concentration.

TABLE 3. NUMBERS OF F. femoralis CAPTURED IN 2 TRAPS, 1 BAITED
WITH SUGAR PLUS (Z)-11-HENTRIACONTENE OR WITH (Z)-ll-
HENTRIACONTENE COMBINED WITH TRICOSANE, AND THE OTHER
WITH SUGAR ALONE.*

Mean number captured
Catch
Amount of test Trap with Trap with ratio:
material treated untreated treated/ Level of
on filter paper filter paper filter paper untreated significance

MALES
5 mg (Z)-ll-C,, 59.7 53.5 1.12 N.S.
25 mg (Z)-ll-C,, 45.0 46.1 0.98 N.S.
5 mg (Z)-ll-C,, plus
5 mg tricosane 51.5 50.6 1.02 N.S.
FEMALES
5 mg (Z)-ll-C3, 35.2 39.5 0.89 N.S.
25 mg (Z)-11-C,, 26.5 34.5 .77 2.5%
5 mg (Z)-ll-C31 plus
5 mg tricosane 43.7 44.4 .98 N.S.

*Means are for fifteen 24-h collections at each concentration.

Catch
ratio:
treated/
untreated

Level of
significance

5%
2.5%
5%

N.S.
N.S.
N.S.

Vol. 61, No. 3, 1978

Uebel et al.: Fannia Mating Stimulants

ACKNOWLEDGMENT
We thank Henry J. Retzer of the Agricultural Equipment Laboratory,
Plant Physiology Institute, FR, SEA, USDA, Beltsville, Maryland, for
designing the rotating platforms used in this study.

LITERATURE CITED
CARLSON, D. A., AND M. BEROZA. 1973. Field evaluations of (Z)-9-tricosene,
a sex attractant pheromone of the house fly. Environ. Ent. 2:555-9.
CARLSON, D. A., M. S. MAYER, D. L. SILHACEK, J. D. JAMES, M. BEROZA,
AND B. A. BIERL. 1971. Sex attractant pheromone of the house fly:
Isolation, identification, and synthesis. Science 174:76-8.
MORGAN, P. B., I. H. GILBERT, AND R. L. FYE. 1974. Evaluation of (Z)-
9-tricosene for attractancy for Musca domestic in the field. Fla. Ent.
57:136-40.
MUHAMMED, S., J. F. BUTLER, AND D. A. CARLSON. 1975. Stable fly sex
attractant and mating pheromones found in female body hydro-
carbons. J. Chem. Ecol. 1:387-98.
ROGOFF, W. M., G. H. GRETZ, M. JACOBSON. AND M. BEROZA. 1973. Con-
firmation of (Z)-9-tricosene as a sex pheromone of the house fly. Ann.
Ent. Soc. Am. 66:739-41.
SONNET, P. E., E. C. UEBEL, R. L. HARRIS, AND R. W. MILLER. 1977. Sex
pheromone of the stable fly: Synthesis and evaluation of methyl-
and 1,5-dimethylalkanes as mating stimulants. J. Chem. Ecol.
3:245-9.
SONNET, P. E., E. C. UEBEL, AND R. W. MILLER. 1975. Sex pheromone of
the face fly and compounds influencing pheromone activity. En-
viron. Ent. 4:761-4.
UEBEL, E. C., M. SCWARZ, R. E. MENZER, AND R. W. MILLER. 1978a. Mating
stimulant pheromone and cuticular lipid constituents of Fannia
pusio (Wiedemann) (Diptera: Muscidae). J. Chem. Ecol. 4:73-81.
UEBEL, E. C., M. SCHWARZ, R. W. MILLER, AND R. E. MENZER. 1978b.
Mating stimulant pheromone and cuticular lipid constituents of
Fannia femoralis (Stein) (Diptera: Muscidae). J. Chem. Ecol. 4:83-93.
UEBEL, E. C., P. E. SONNET, B. A. BIERL, AND R. W. MILLER. 1975a. Sex
pheromone of the stable fly: Isolation and preliminary identifica-
tion of compounds that induce mating strike behavior. J. Chem. Ecol.
1:377-85.
UEBEL, E. C., P. E. SONNET, R. E. MENZER, R. W. MILLER, AND W. R.
LUSBY. 1977. Mating stimulant pheromone and cuticular lipid con-
stituents of the little house fly, Fannia canicularis (L.). J. Chem.
Ecol. 3:269-78.
UEBEL, E. C., P. E. SONNET, AND R. W. MILLER. 1976. House fly sex phero-
mone: Enhancement of mating strike activity by combination of (Z)-
9-tricosene with branched saturated hydrocarbons. Environ. Ent.
5:905-8.
UEBEL, E. C., P. E., SONNET, R. W. MILLER, AND M. BEROZA. 1975b. Sex
pheromone of the face fly, Musca autumnalis De Geer (Diptera:
Muscidae). J. Chem. Ecol. 1:195-202.

The Florida Entomologist

BOOK REVIEW

DEFENSE MECHANISMS OF PLANTS. Brian J. Deverall. 1977.
Cambridge University Press: Cambridge, England. 110 p. $12.50. This is the
19th monograph in the series, Cambridge Monographs in Experimental Bi-
ology. The author is Professor of Plant Pathology at the University of
Sydney.
The book is devoted entirely to plant defense against fungal and bac-
terial plant pathogens. By Professor Deverall's definition, a defense mech-
anism is "the dynamic processes by which plant cells perceive the approach
of an intruder and occasionally permit, but usually discourage, its further
progress." The title of the book implies that more than 1 defense mecha-
nism exists in plants to pathogens, but in reality, only 1 such mechanism,
hypersensitivity, is discussed. This defense mechanism seems to be present in
all plants, but expression of it is controlled by interactions of single genes
in the plant and the pathogen.
Professor Deverall discusses the hypersensitive reaction as a dynamic
process and is correct in doing so. Hypersensitivity includes: (1) ingress of
pathogen, (2) initial growth of pathogen, (3) induction of the reaction, (4) re-
sponse of plant cell, (5) plant cell collapse and necrosis, and (6) forma-
tion of antifungal, or antibacterial, compounds (phytoalexins). Each of
these stages are mentioned by Professor Deverall, but most emphasis is on
the last stage. Thirty percent of the book concerns the induced formation
and biosynthesis of phytoalexins, and the role of phytoalexins in the de-
fense mechanism. It is obvious that Professor Deverall considers phyto-
alexins responsible for cessation of growth of pathogens in the hypersensi-
tive reaction. Not everyone in research on hypersensitivity agrees with this
position. Some of the papers with the different point of view are mentioned,
but not discussed thoroughly.
The hypersensitive reaction is one of the major mechanisms for plant
resistance to disease. It is a very effective type of resistance, conferring "field
immunity." Hypersensitivity occurs not only to plant pathogenic fungi and
bacteria, but to plant viruses, and nematodes as well. This book will give
everyone interested in plant resistance to pathogens an up-to-date review
of the physiology of this important resistance mechanism.
The book has an extensive list of references. These references are inval-
uable to teachers and students of plant pathology. It also illustrates the
interest and research that has occurred in recent years on this very important
resistance mechanism.

Robert E. Stall
Department of Plant Pathology
University of Florida
Gainesville, FL, 32611

Vol. 61, No. 3, 1978

The Florida Entomologist

PHYSICAL AND BIOLOGICAL EVALUATIONS
OF AN ENCAPSULATED DIET FOR REARING
CHRYSOPA CARNEA1,23

P. B. MARTIN,4 R. L. RIDGWAY,5 AND C. E. SCHUETZE6
ABSTRACT
Evaluations were made of an encapsulated diet developed for rearing
the green lacewing, Chrysopa carnea Stephens. Sizes of the capsules were
similar to those of eggs of tobacco budworm, Heliothis virescens (F.), but
considerably larger than those of eggs of Angoumois grain moth, Sitotroga
cerealella (Olivier). Capsules were more resistant to dehydration than bud-
worm eggs, but less resistant than grain moth eggs. Few Ist-instar larvae of
C. carnea penetrated capsules; however, 2nd- and 3rd- instar larvae had
little difficulty. All stages were effective in penetrating the shells of insect
eggs. Some stages of Geocoris punctipes (Say), Reduviolus sp., Orius in-
sidiosus (Say), Zelus renardii Kolenati, Coleomegilla maculata DeGeer,
Hippodamia convergens Guerin-M6neville, and Collops sp. also fed on en-
capsulated diet.

Considerable effort has been made to develop an artificial diet for
efficiently rearing Chrysopa carnea Stephens (Hagen and Tassan 1965,
Ridgway and Kinzer 1974). Vanderzant (1969) reported a semi-defined diet
(presented in cellulose sponge) for rearing larvae of C. carnea. This diet
was subsequently improved (Vanderzant 1973); however, a practical method
of feeding the diet to large numbers of insects was needed. After evaluation
of about 50 different materials, a capsule of paraffin wax, candelilla wax,
polyethylene, and polybutene was selected (Anonymous 1971). These com-
ponents prevented losses of substantial amounts of diet and yielded a
capsule with physical characteristics which could be punctured by larvae
of C. carnea. Subsequent tests showed that the encapsulated diet could be
used in mass-rearing several generations of C. carnea (Ridgway et al. un-
published report).'
Insect development was slower and mortality was higher among C.
carnae larvae reared on capsules than on eggs of the Angoumois grain
moth, Sitotroga cerealella (Olivier) (Ridgway et al. unpublished report).7
Eggs of Heliothis spp. also were apparently preferred over capsules by
larvae of C. carnea. Obviously, some improvements in the capsule are
needed to provide for more reliable production of C. carnea.

Neuroptera: Chrysopidae.
This research was supported in part by Cotton, Inc., 4505 Creedmoor St., Raleigh, N.C.
27612. Received for publication 4 October 1977.
'This paper is a portion of a thesis submitted by the senior author to Texas A & M University
in partial fulfillment of the requirements of the M.S. degree.
SFormerly, Texas A & M University, College Station, Texas; now Dep. Entomol.-Fisheries,
Georgia Coastal Plain Experiment Station, Tifton, Ga. 31794.
'Formerly, Cotton Insects Research Laboratory, ARS, USDA, College Station, Texas;
now National Program Staff, SEA, USDA, Beltsville, Md. 20705.
6 Southwest Research Institute, San Antonio, Texas 78200.
7Ridgway, R. L., V. S. House, and E. S. Vanderzant. 1969. Evaluation of encapsulated
diets for rearing Chrysopa. Annu. Rep. (Ent. Res. Div., Agric. Res. Serv., USDA, College
Station, Texas).

Vol. 61, No. 3, 1978

146 The Florida Entomologist Vol. 61, No. 3, 1978

To develop information of value in improving capsules for mass-rearing
C. carnea, we compared some physical properties of the capsules with eggs
of the tobacco budworm, Heliothis virescens (F.), and the Angoumois grain
moth. Also, some feeding characteristics of C. carnea and several other pre-
daceous arthropods were compared on encapsulated diet and on budworm
and grain moth eggs.

MATERIALS AND METHODS
Experiments proceeded under constant light at 27+2C. Tobacco bud-
worm and Angoumois grain moth eggs usually less than 24-hr old were
collected from laboratory cultures.
The capsules had a shell of paraffin wax (80%), candelilla wax (10%),
polyethylene (5%) and polybutene (5%). They were made with an experi-
mental device developed at Southwest Research Institute, San Antonio,
TX (Anonymous 1971). The diet encapsulated was similar to that reported
by Vanderzant (1969) except that it contained blue dye (0.04% F. D. and
C. Blue Dye No. 1).
PHYSICAL PROPERTIES.-We measured the diameter and shell thickness of
capsules and eggs (maximum length and diameter of the elongate grain
moth eggs) with a ocular micrometer on a phase-contrast microscope (100
and 400 X). Capsules were split in half with teasing needles on glass slides;
measurements were made on 3 samples of 25 eggs or capsules. In addition,
weights were determined with an analytical balance by using 3 samples of
25 to 200 eggs or capsules.
We determined fill (liquid contents by weight) of the capsules and eggs
by obtaining a gross weight of 3 to six 250-350 mg samples and subtracting
the weight of shell material. The shell was isolated by crushing the cap-
sules or eggs on filter paper and rinsing the liquid diet through the filter
with distilled water. The paper-shell combination was then dried and
weighed. Initial weight of the filter paper was subtracted to obtain weight
of shell material. Another filter paper of the same weight was saturated
with water, dried, and used as a control. Percent fill (by weight) was cal-
culated by dividing the weight of contents by the gross weight (% fill = 100
(fill-wt)/(gross-wt)). Approximate volume held in individual capsules or
eggs was calculated by dividing the weight of the contents by their estimated
density and multiplying by percent fill. (Contained-vol = (wt/d) x %
fill.) To determine density, a sample of capsules or eggs (ca. 0.5 g) was
weighed and allowed to sink in a 5 ml graduated cylinder half full of
water. Volume displaced was measured and divided into sample weight.
Loss of fill (dehydration) from capsules, tobacco budworm eggs, and
Angoumois grain moth eggs was determined at 3 different relative humidities
(RH) over a 2- or 3-day period. Capsules were taken from refrigeration,
held at room temperature for 30-45 min, weighed, placed on small, open
plastic containers (5 cm in diam), arid held in a controlled RH system.
Initial weight (200-300 mg) and changes in weight of capsules were measured
on an analytical balance. Tests on the capsules, budworm eggs, and grain
moth eggs were replicated 6, 9, and 12 times, respectively. Samples were
held in airtight glass humidity chambers containing saturated solutions
(Winston and Bates 1960) of either lithium chloride (ca. 12% RH), potassium
nitrate (ca. 48% RH), or potassium sulfate (ca. 97% RH). Since the humidity

Martin et al.: Encapsulated Diet for Chrysopa

chambers were opened to room conditions when weighing capsule samples,
the 3 different RH will be referred to as low, medium, and high in the dis-
cussion below.
The number of eggs (tobacco budworm and Angoumois grain moth)
whose fill was equal to that of a mean capsule (capsule-equivalents)
was estimated from their respective volumes (which had been adjusted for
mean loss of fill after 48 hr at about 48% RH). We calculated this by
dividing mean adjusted capsule volume by mean adjusted budworm and
grain moth egg volumes. (Cap-equiv = (cap-vol X % loss) / (egg-vol X
% loss).)
We examined three lots of capsules with a dissecting scope (50 X)
for deviations from the desired uniform, spherical capsule. Four samples
of 100 capsules from each lot were examined for capsules containing holes
and lobes, as well as shell material in their interior. Several other lots
were examined for capsules containing holes greater than 30 p in diameter.
Loss of fill of the different lots was measured at a low RH (ca. 12%).
COMPARATIVE FEEDING CHARACTERISTICS.-We compared encapsulated
diet with insect eggs by "bioassays" with C. carnea and several other
predaceous arthropods. Preferences, consumption, and penetration effi-
ciency were determined for all 3 instars of C. carnea larvae although we
concentrated on the more delicate Ist-instar larvae.
Larvae of C. carnea used in these studies were taken from a laboratory
culture that had been reared solely on Angoumois grain moth eggs prior
to testing. Rearing techniques were similar to those described by Ridgway
et al. (1970).
Chrysopa carnea Preference and Consumption.-Capsules and eggs (either
budworm or grain moth) were placed together and offered to individual
larvae (all 3 instars). Various combinations of eggs and capsules were
placed at random in the bottom of 50 x 12 mm plastic petri dishes that
contained a single larvae each. Consumption of eggs and encapsulated diet
were recorded after 24 hr. First-instar individuals were unfed, starved 0-6 hr
and confined with 50-125 eggs or capsules (replicated 10 and 12 times8);
2nd-instar larvae were starved 6-10 hr and offered 50-125 eggs or capsules
(replicated 10 and 19 times"); and 3rd-instar larvae were starved 6-12 hr
and offered 100-125 eggs or capsules (replicated 5 and 17 times8).
In a 2nd experiment, consumption of encapsulated diet, tobacco bud-
worm and Angoumois grain moth eggs was determined by the same procedure
(replicated 13 17 times) except that only one food source was offered each
larva. To maximize consumption, at least 400 eggs or capsules were offered
to each individual.
In addition, relative consumption of capsules and eggs was measured by
weight gain. Unfed Ist-instar larvae of C. carnea were confined in 15-ml
plastic cups containing excessive amounts of encapsulated diet, tobacco
budworm eggs, or Angoumois grain moth eggs. After 48 hr, we weighed the
larvae to determine the amount of food consumed and recorded percent
mortality. Thirty-five to 48 larvae were tested in each of the 3 groups.
Chrysopa carnea Penetration Efficiency.-Larvae of each instar of C. carnea
were placed individually in 100 x 15 mm petri dishes containing ca. 500
budworm or grain moth eggs, or capsules. First-instar larvae (8 replicates)

I For grain moth egg-capsule and budworm egg-capsule combinations, respectively.

The Florida Entomologist

were observed for 30 min while 2nd- and 3rd-instar larvae (9 10 replicates)
were observed for 10 min. Counts were made of the number of times larvae
made definite contact with capsules (or eggs) with their mandibles versus
the times they penetrated the food source. Penetration efficiency was then
determined by calculating the mean percent of total contacts that were
followed by penetration.
We also determined if Ist-instar larvae which are successfully cultured
on encapsulated diet might be obtaining diet by feeding through holes in
incomplete capsules (structurally defective). Single eggs of C. carnea
nearing eclosion were confined with complete capsules, incomplete cap-
sules, or Angoumois grain moth eggs. We offered each insect 10 eggs or
10 capsules. Forty separate eggs of C. carnea were placed with each food
source in depressions (3 mm deep X 8 mm diam) in a steel plate. The steel
plate was then covered with plate glass. Twenty-four to 48 hr after eclo-
sion, larvae of C. carnea were observed for evidence of feeding. Larval
mortality was also recorded.
Feeding by Other Predaceous Arthropods.-Adults and immature stages of
Geocoris punctipes (Say), Reduviolus sp., Zelus renardii Kolenati, and
Coleomegilla maculata DeGeer; immature stages of Orius insidiosus (Say)
and Hippodamia convergens Gu6rin-M6neville; and adults of Collops sp.,
were used for additional similar evaluations of capsules. The insects were
observed individually for feeding behavior, survival, and development.

RESULTS
PHYSICAL PROPERTIES.-Physical measurements indicated that capsules
were similar to tobacco budworm eggs in size, weight, and volume of fill
contained (Table 1). Well-formed capsules were near-perfect spheres,
tobacco budworm eggs were somewhat oval, and Angoumois grain moth eggs
were oblong and cigar-shaped. The shell textures of tobacco budworm and
Angoumois grain moth eggs were rough and ridged, while the capsule was
generally smooth and slick. The mean diameter of tobacco budworm
TABLE 1. COMPARATIVE PHYSICAL PROPERTIES OF ENCAPSULATED Chry-
sopa carnea DIET, AND EGGS OF THE TOBACCO BUDWORM AND
THE ANGOUMOIS GRAIN MOTH.

Food Source
Properties measured Capsules Budworm eggs Grain moth eggs

Size (max. outside 458 + 58 555 + 23 251 + 20 X 625 + 36
diam in fL)*
Shell thickness (t)* 13 + 5 45 4
% fill** 82.0 + 0.3 94.9 + 1.2 93.4 + 1.0
Weight in tg t 56 + 1.0 78 5.7 10 + 0.7
Volumein cm3tt 4.2 X 10-5 6.7 x 10-1 1.7 x 10-5
Capsule-equivalents --- 0.7 2.0

*Mean + SD from 3 samples of 25 each.
**Mean + SD from 3-6 samples of 250-350 mg each.
t Mean + SD from 3 samples of 25-200 eggs or capsules.
ft Estimated from weight, ca. specific gravity (1.1 g/cm), and % fill.

Vol. 61, No. 3, 1978

Martin et al.: Encapsulated Diet for Chrysopa

eggs was greater than that of capsules; Angoumois grain moth eggs were
longer than capsules but their diameter was smaller. Insect eggs were more
uniform in size than capsules. The shell of the capsules was at least 2.6
times thicker than eggs from either species of insect. Capsules contained a
lower percentage of liquid contents than eggs, but actually possessed more
liquid content than did individual Angoumois grain moth eggs.
At all humidities the most fill was lost by tobacco budworm eggs
followed by capsules and Angoumois grain moth eggs (Fig. 1). After 48 hr
at the high RH, loss of fill was <15% for the 3 food sources tested; at
the medium relative humidity, loss of fill averaged 42% for tobacco bud-
worm eggs, 26% for capsules, and 9% for Angoumois grain moth eggs; and
at the low RH, loss of fill was 59% for tobacco budworm eggs, 28% for
capsules, and 14% for Angoumois grain moth eggs.
Variation in frequency of incomplete capsules and in sizes of the holes in
the capsules (structural defects) apparently accounted for most of the var-
iation in loss of capsule fill. Capsules from the lot which contained the

40

= 30

20

10

40

6 30 4

20
= Xso ,.'
10

50

H 12 24 36 48
HOURS
Fig. 1. Effect of different relative humidities on loss of fill
carnea encapsulated diet, tobacco budworm (Heliothis)
goumois grain moth (Sitotroga) eggs.

from Chrysopa
eggs, and An-

The Florida Entomologist

smallest number of holes (3.8%) larger than 30 t lost the least amount of
fill (9.0%) after 48 hr, while the lot with the highest percent of large holes
(29.8%) lost the most fill after 48 hr (25.3%). An increase in frequency of
incomplete capsules was positively correlated with an increase in fill of
the capsules and decrease in the shell thickness. Capsules with odd shapes,
multiple lobes, and shell material in the interior occurred in nearly
equal frequencies among the 3 lots evaluated (11.2, 12.5, and 12.8%).
COMPARATIVE FEEDING CHARACTERISTICS.-Preference and Consumption.-
All 3 instars of C. carnea larvae, confined individually with tobacco bud-
worm eggs and encapsulated diet, fed on more and consumed more content
from budworm eggs than capsules (Table 2). First- and 2nd-instar larvae
fed on more and consumed more contents of Angoumois grain moth eggs
than that of encapsulated diet whereas, 3rd-instar larvae consumed about
equal amounts of capsules and capsule-equivalents of Angoumois grain
moth eggs. When offered separately, no significant differences in volumes of
encapsulated diet, budworm eggs, and grain moth eggs were consumed by
2nd- and 3rd-instar larvae. First-instar larvae consumed significantly less
volume from capsules than from eggs of either budworm or grain moth.
Insects that died were not included in calculating consumption values.
A higher percent mortality (32%) occurred among those Ist-instar larvae
of C. carnea offered only encapsulated diet than among those offered
insect eggs (15%). No mortality occurred among 2nd- or 3rd-instar larvae
regardless of the food source. First-instar larvae of C. carnea surviving
after 48 hr on encapsulated diet gained approximately 135 ug. Those surviv-
ing on budworm and grain moth eggs gained 452 and 345 tg, respectively,
during the same period of time.

TABLE 2. COMPARATIVE 24-HR CONSUMPTION OF ENCAPSULATED DIET,
TOBACCO BUDWORM EGGS, AND ANGOUMOIS GRAIN MOTH EGGS
BY Chrysopa carnea LARVAE.

Mean actual no. consumed Mean capsule-equivalents
by indicated instar* consumed by indicated instar**

Food source First Second Third First Second Third

OFFERED IN COMBINATION
Capsules vs. 0.1 0.9 14.4 0.la 0.9a 14.4a
Budworm eggs 1.5 5.2 36.3 2.1b 7.4b 51.4b
Capsules vs. 0.2 0.4 64.2 0.2a 0.4a 64.2a
Grain moth eggs 6.3 9.7 123.0 3.1b 4.8b 61.5a
OFFERED SEPARATELY
Capsules 1.4 54.6 200.0 1.4x 54.6x 200.0x
Budworm eggs 4.7 27.5 110.7' 6.7y 39.3x 157.3x
Grain moth eggs 8.4 95.4 301.0 4.2z 47.8x 150.8x
*Mean no. consumed by 5-19 larvae in individual petri dishes containing food source.
**No. of eggs equivalent to the mean fill of capsules; calculated from respective volumes
that had been adjusted for mean loss of fill after 48 hr at ca. 48% RH.
Sets of numbers listed vertically with the same letter (a or b for the "combination" sets,
or x, y, or z for the "separate" sets) are not significantly different according to Student's t test
for paired observations (combination), or Duncan's multiple range test (separate), at the 5%
level.

Vol. 61, No. 3, 1978

Martin et al.: Encapsulated Diet for Chrysopa

Penetration Efficiency.-All 3 larval instars of C. carnea were less efficient
in piercing capsules than eggs (Table 3). First-instar larvae penetrated a
mean of 46% of the Angoumois grain moth eggs and 14% of the tobacco
budworm eggs, but no Ist-instar larvae were observed to penetrate the cap-
sules in the 30 min periods. The penetration efficiency of 2nd- and 3rd-
instar larvae was significantly higher on eggs than on capsules. No distinct
attack phase in the feeding process was noticed for larvae observed in these
penetration efficiency tests. Usually, the first definite indication of pene-
tration was feeding action by the larvae or the beginning of collapse of
an egg. When capsules were penetrated, the mandible usually could be
seen easily through the shell.
In the 2nd test individual larvae were left with the capsules for 24-48
hr, and had not been fed previously. This time Ist-instar larvae of C. carnea
did penetrate the capsule but again they used Angoumois grain moth eggs
much more effectively than capsules. Complete capsules were fed upon by
30.5% of the larvae. But 38.2% fed on incomplete capsules and 54.5% fed on
Angoumois grain moth eggs.
Feeding on Encapsulated Diet by Other Predaceous Arthropods.-Some
stages of all of the other predators studied fed on the encapsulated diet,
but it appeared to be more suitable for Reduviolis sp. and C. maculata. We
reared both of these species from 2nd-instar to adult on encapsulated
diet. Early instars of these predators generally had difficulty in utilizing
the encapsulated diet, apparently because of difficulty in penetrating the
capsule.

TABLE 3. PENETRATION EFFICIENCY OF Chrysopa carnea LARVAE ON
CAPSULES, AND ON EGGS OF THE TOBACCO BUDWORM AND AN-
GOUMOIS GRAIN MOTH.

Ratio of total Mean penetration
penetrations to efficiency/insect
Food source contacts by instar* (%)**

FIRST INSTARt
Capsules 0/1215 Oa
Budworm eggs 4/422 14b
Grain moth eggs 17/179 46c
SECOND INSTAR tt
Capsules 60/211 58a
Budworm eggs 36/40 94b
Grain moth eggs 105/109 97b
THIRD INSTAR ft
Capsules 191/266 76a
Budworm eggs 95/97 99b
Grain moth eggs 220/232 95b

*Total contacts and penetrations.
**Mean percent over 8-10 replicates. Numbers followed by the same letter are not significantly
different according to Duncan's multiple range test at the 5% level.
t Eight insects observed for 30 min.
t t Nine or 10 insects observed for 10 min.

152 The Florida Entomologist Vol. 61, No. 3, 1978

DISCUSSION
The use of high humidities to reduce loss of fill from capsules should
result in maximum utilization of encapsulated diet without causing rearing
problems. Finney (1948) reported 80% RH was desirable for rearing larvae
of C. carnea. High humidity should also be favorable to the successful
utilization of the encapsulated diet.
Although the Angoumois grain moth-reared C. carnea used in these
studies had some difficulty in adapting to the encapsulated diet, there are
indications that a more successful strain can be selected (Ridgway et al.
unpublished report9). Grain moth eggs could be used to supplement the
capsules in the Chrysopa diet. This does not negate the need for improve-
ment of the capsules. Data from this study indicate that difficulty of
Ist-instar C. carnea (and other predaceous arthropods) in feeding on the
encapsulated diet may be an important problem to overcome for successful
implementation of the capsule in mass production of C. carnea.

LITERATURE CITED
ANONYMOUS. 1971. Package meals for insects. Agric. Res. USDA. 19:3-4.
FINNEY, G. L. 1948. Culturing Chrysopa californica and obtaining eggs for
field distribution. J. Econ. Ent. 41:719-21.
HAGEN, K. S., AND R. L. TASSAN. 1965. A method of providing artificial
diets to Chrysopa larvae. J. Econ. Ent. 58:999-1000.
RIDGWAY, R. L., AND R. E. KINZER. 1974. Chrysopids as predators of crop
pests. Entomophaga 7:45-51.
RIDGWAY, R. L., R. K. MORRISON, AND M. BADGLEY. 1970. Mass rearing a
green lacewing. J. Econ. Ent. 63:834-6.
VANDERZANT, E. S. 1969. An artificial diet and a rearing method for Chry-
sopa carnea larvae and adults. J. Econ. Ent. 62:256-7.
VANDERZANT, E. S. 1973. Improvements in the rearing diet for Chrysopa
carnea and the amino acid requirements for growth. J. Econ. Ent.
66:336-8.
WINSTON, P. R., AND D. H. BATES. 1960. Saturated solutions for the control
of humidity in biological research. Ecology 41:232-7.

Ridgway, R. L., R. E. Kinzer, and S. L. Jones. 1971. Ability of larvae from two laboratory
strains of Chrysopa to consume an encapsulated diet and Sitotroga eggs. 2nd Quarterly Rep.
(Ent. Res. Div., Agric. Res. Serv., USDA, College Station, Texas).

The Florida Entomologist

USING FROZEN HOST PUPAE' TO INCREASE THE
EFFICIENCY OF A PARASITE2-RELEASE PROGRAM3

L. G. PICKENS, AND R. W. MILLER

Chemical and Biophysical Control Laboratory
Agricultural Environmental Quality Institute
Federal Research, Science and Education Administration
USDA, Beltsville, Md. 20705

ABSTRACT
Continued, periodic additions of frozen house fly pupae, Musca domes-
tica L., were used to augment an adult-release program of the wasp Pachy-
crepoideus vindemiae (Rondani) in chicken houses in Maryland. A one-
month long wasp release program, which was supplemented and followed
by deposits of frozen fly pupae, was equally or more effective in killing
house fly pupae than a program of continued releases of adult wasps.
Both programs resulted in an increased house fly pupal mortality of 22-44%
compared to that in an untreated check house.

Considerable research has been done to develop a biological control
program for flies in chicken houses (Axtell 1970, Legner and Brydon 1966,
Morgan et al. 1975, Pickens et al. 1975). House flies, Musca domestic L.,
and little house flies, Fannia canicularis (L.), breed in accumulated chicken
feces in commercial poultry houses. The manure is often difficult to com-
pletely remove or to treat with insecticides, but it does remain in a fixed
and well-defined area which facilitates some parasite manipulations.
One limitation of previously developed parasite-release programs has
been the need to make continued, regularly timed releases of parasites
in each house in order to compensate for parasite mortality and emigration
in periods when few host insects are present. A supplemental method of
maintaining numbers of parasites or of increasing their hunting efficiency
in periods of low host density would be to artificially add suitable stages
of the host to the environment in a condition which would prevent them
from developing to the pest stage of their life cycle if they should fail
to be parasitized.
Since the wasp Pachycrepoideus vindemiae (Rondani) will parasitize
dead house fly pupae (Crandall 1939), and is an active hunter of fly pupae
in chicken feces (Pickens et al. 1975), we used them to evaluate the feasi-
bility of using dead 2-day-old house fly pupae to improve the efficiency
of an adult wasp release program against flies breeding in chicken houses.
We compared fly pupal mortality and parasite production in chicken
houses maintained only with regular releases of parasites versus houses
initially maintained by parasite releases but regularly supplemented by
deposits of dead host pupae; a third house that received neither parasites
nor dead host pupae served as a check. Observations were also made on

SDiptera: Muscidae.
2 Hymenoptera: Pteromalidae.
SReceived for publication 17 February 1978.

Vol. 61, No. 3, 1978

The Florida Entomologist

pest mortality due to parasites when the parasite release and pupal houses
were relatively close together (30m) as opposed to farther apart (100m).

MATERIALS AND METHODS
In 1975, we released ca. 1000 adult P. vindemiae wasps per m2 of feces
3 times a week from 12 May to 12 Sept. in one house, and 1000 wasps per
m2 of feces from 12 May to 12 June, plus ca. 5000 frozen 2-day-old house
fly pupae every Monday, Wednesday, and Friday from 12 May to 12 Sept.
The pupae were frozen when they were 2 days old and were acceptable
to the wasps for ca. 48 h after thawing. A third "check" house received
neither wasps nor frozen pupae. The houses used in the tests measured ca.
4 x 4 x 3 m high, were open on the south side, and each housed 24 chickens
in a wire cage suspended 1 m above the floor. Feces were allowed to
accumulate beneath the cages during the test. All tests were conducted
on the USDA Agricultural Research Center, Beltsville, MD.
In 1976, 300 P. vindemiae wasps per m2 of feces were released every
Monday and Wednesday in each of two houses from 17 May to 23 June. One
of the houses received only the wasps, while the other house received wasps
plus ca. 3000 frozen house fly pupae each Monday and Wednesday from
17 May to 1 Sept. A third "check" house received neither wasps nor frozen
pupae. Fewer wasps were released in 1976 than in 1975 because of problems
with the colony.
In 1975, the treated houses were ca. 30 m apart in a north-south line, and
100 m east of the check house. In 1976, they were ca. 100 m apart and 1 km
east of the check house. Prevailing winds were from the southwest in both
years.
All of the released wasps were 1-day-old adults; they were released
on the floor along the western edge of the fecal packs in each house. The
frozen pupae were also placed along the western edge of the fecal pack.
The effect of the treatments was determined by collecting up to 100 of
the pupae of each fly species from the surface and the first 5 cm depth of
the eastern edge of each fecal pack with forceps every Monday, Wednes-
day and Friday during the tests. The three species of fly pupae collected
were M. domestic, F. canicularis, and the predatory black garbage fly,
Ophyra leucostoma (Wiedemann). The collected pupae were held in con-
tainers in the laboratory for 30 days at 270C and then checked for mortality
and/or wasp eclosion and species. The effectiveness of a treatment was
expressed as both the number and percentage of the weekly pupal sample
that produced adult wasps or that died (compared to the percentages for
similar samples from the check houses). The fraction of frozen pupae which
had been parasitized in 2 days was determined by collecting pupae from the
frozen pupae piles of the prior deposit.
All count means were compared by an ANOVA and Duncan's new mul-
tiple range test with significance equal to P 40.05. Each week was con-
sidered a replicate because treatments were the same for all weeks within
each year and because each generation of fly pupae lasted ca. 1 week.

RESULTS AND DISCUSSION
In both years, the greatest number of house fly pupae were parasitized

Vol. 61, No. 3, 1978

Pickens and Miller: Using Frozen Host Pupae

by P. vindemiae in the house to which frozen pupae were added; the smallest
number were parasitized by P. vindemiae in the check houses, which received
neither wasps nor frozen pupae (781 and 255 vs 484 and 58 in 1975 and 1976,
respectively). In 1975, there was little difference between the total number
of fly pupae collected over the summer from any of the 3 houses (2246,
2415, and 2582) but in 1976 the greatest number were collected from the
check house and the least from the frozen pupal house and the wasp release
house (2971 vs 1831 and 1622). Approximately equal percentages of fly
pupae were parasitized in both treated houses in both years, and a signifi-
cantly lower percentage were parasitized in the untreated house in both
years, except for F. canicularis in 1975 (Table 1). No parasites except P.
vindemiae emerged from any of the collected pupae.
The number of pupae available for parasitization and the percentage
of pupae which produced wasps both declined throughout 1975 indicating
that the rate of parasitization was directly related to host pupal density,
but only the percentage parasitized declined in 1976 (Fig. 1), which suggests
that the rate of parasitization in 1976 was limited by the low numbers
of wasps present. The wasps parasitized approximately equal percentages
of frozen vs living pupae (20% vs 30% and 10% vs 9% in 1975 and 1976,
respectively). Although the house which received wasps plus frozen pupae
had a more or less constant percentage of parasitization in the first 12 weeks
of 1975, the house which received only wasps did not have a constant per-
1975
120
WASPS FROZEN PUPAE C CHECK

80
60 4

40 A -

0 A A

o-b No. of pupae collected (+3) a--A% parasitized

1976
120
WASPS FROZEN PUPAE CHECK
100
80
60 -
40 -
20

2 4 6 8 10 12 14 2 4 6 8 10 12 14 2 4 6 8 10 12 14
A A a
WEEKS AFTER FIRST WASP RELEASE
Fig. 1. The total number of pupae of 3 fly species, Musca domestic
L., Fannia canicularis (L.) and Ophyra leucostoma (Weidemann), which
were collected and the percentages of them which were parasitized by
Pachycrepoideus vindemiae in chicken houses. ( A indicates the end of
adult releases, A on the x-axis indicates the end of frozen house fly pupal
additions.)

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Vol. 61, No. 3, 1978

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Pickens and Miller: Using Frozen Host Pupae

centage of parasitized pupae beyond the 7th week after the initial wasp re-
lease in 1975. Neither method maintained a constant number or percentage
of pupae parasitized beyond the 7th week in 1976. The house which received
frozen pupae had a higher second peak of parasitization than did the house
which received only adult wasps (in the 10th week in 1975 and in the 7th
week in 1976).
Large numbers of wasps apparently dispersed to at least 100 m in 1975,
since there were many fly pupae parasitized at the check house after the
second week of the tests. Very few pupae were parasitized at the more distant
(1 km) check house in 1976 (Table 1).
The percentage mortality was significantly greater in the treated houses
than in the check house in both years (Table 1), indicating that much of the
mortality was due to the wasps. Since P. vindemiae adult females kill as
many pupae for food as they use for oviposition (Pickens et al. 1975), this
is consistent with the wasp's behavior.
There was a close direct relationship between changes in the number of
fly pupae present and in the number parasitized, but no relationship between
the number parasitized and the numbers of wasps released. This would
suggest that the wasps tended to remain in or to hunt longest in areas with
large numbers of pupae (Legner 1967).
Since there was a rapid (2-day) decline in the number of pupae which
were parasitized after each peak of parasitization and a close correlation
between the number of pupae present and the number parasitized in 1975,
the wasps apparently dispersed rapidly from their emergence sites and were
then attracted to areas where pupae were concentrated (Pickens et al. 1975).
This would also explain why the addition of frozen pupae resulted in in-
creased parasitization when placed within 50 m of a wasp release site; i.e.,
the wasps dispersed from emergence or release sites, but remained near
sites of pupal concentration. In our earlier studies (Pickens et al. 1975), we
observed that pupae which were 4 m distant from a wasp emergence site
in a dark building were not parasitized, even though large numbers of
female wasps went to the building windows and to U.V. lamps. Nostvik
(1954) also reported that the female wasps are strongly attracted to light.
Neither the continued periodic releases of adult wasps nor the con-
tinued periodic additions of frozen host pupae maintained a constant wasp
population in either chicken house in the absence of immigration from
another wasp release site. The use of frozen pupae in houses either 30 m or
100 m from a wasp release site apparently improved the efficiency of
parasitization in the pupae houses; parasitization of living fly pupae was
generally higher than that observed for houses receiving only wasp re-
leases. The dead fly pupae apparently either attracted or delayed searching
wasps. The data suggest that dead host pupae may be manipulated to in-
crease the efficiency of a wasp release program; 2 houses (one wasp re-
lease, one pupae house) were comparably parasitized by the same number of
wasps required for release in one house if only adult wasps were used.
Although the manipulation of P. vindemiae did result in an increased
house fly pupal mortality of 22-44% over that in the check houses, this
parasite is a relatively poor searcher compared to other species of wasps
and is probably best used in programs where fly breeding is concentrated
in a small area.

The Florida Entomologist

LITERATURE CITED
AXTELL, R. C. 1970. Integrated fly-control program for caged-poultry
houses. J. Econ. Ent. 63:400-5.
CRANDALL, H. A. 1939. The biology of Pachycrepoideus dubious Ashmead
(Hymenoptera), a pteromalid parasite of Piophila casei L. (Diptera).
Ann. Ent. Soc. Am. 32:632-51.
LEGNER, E. F., AND H. W. BRYDON. 1966. Suppression of dung-inhabiting
fly populations by pupal parasites. Ann. Ent. Soc. Am. 59:638-51.
LEGNER, E. F. 1967. Behavior changes in the reproduction of Spalangia
cameroni, S. endius, Muscidifurax raptor, and Nasonia vitripennis
(Hymenoptera: Pteromalidae) at increasing fly host densities. Ibid.
60:819-26.
MORGAN, P. B., R. S. PATTERSON, G. C. LABRECQUE, D. E. WEIDHAAS,
AND A. BENTON. 1975. Suppression of a field population of house
flies with Spalangia endius. Science 189:388-9.
NOSTVIK, E. 1954. Biological studies of Pachycrepoideus dubious Ashmead
(Chalcidoidea: Pteromalidae), a pupal parasite of various Diptera.
Oikos 5:195-204.
PICKENS, L. G., R. W. MILLER, AND M. M. CENTALA. 1975. The biology, pop-
ulation dynamics, and host finding efficiency of Pachycrepoideus
vindemiae in a box stall and a poultry house. Environ. Ent. 4:975-9.

SSTORTER

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Vol. 61, No. 3, 1978

The Florida Entomologist

ECOLOGY AND TAXONOMY OF LOWER
RIO GRANDE VALLEY ZETHUS12
CHARLES C. PORTER"

Department of Biological Sciences
Fordham University, Bronx, N.Y. 10458
ABSTRACT
Lower Rio Grande Valley populations of Zethus miscogaster Saussure,
Z. montezuma Saussure, Z. otomitus Saussure, and Z. aztecus Saussure
vary dramatically in abundance from year to year and within each year
show some differences in monthly phaenology. They are active well into
winter and fly on any day when the shade temperature reaches 22C. All
visit the same rather wide range of flowers. They are Neotropic taxa which
approach their northern limit in Texas. Z. otomitus is recorded for the 1st
time from the United States.

In a review of the Lower Rio Grande Valley Zethus (Porter 1975b),
I reported Z. aztecus and Z. montezuma as new for the United States, cited
Z. miscogaster for the 1st time from Texas, and offered a key to all United
States members of this primarily Neotropic genus. After more collecting in
Hidalgo County, Texas, at the Bentsen Rio Grande Valley State Park and
the Valley Botanical Garden, I have obtained more specimens and eco-
logical data on miscogaster, montezuma, and aztecus. I have established
also the presence in south Texas of Z. otomitus, a species hitherto unknown
north of central M6xico.
Zethus are strong-flying vespoids that provision their nests, made in
twigs or trunks or attached to twigs, with lepidopterous larvae. Of the
187 American species, 98 either enter or are restricted to the evergreen and
deciduous forests of tropical South America (Bohart and Stange 1965:22).
Middle America has 36 species, and all throughout its range Zethus tends
to evolve forms adapted to thorn scrub or even deserts. It is thus not sur-
prising that 4 species have reached the semiarid Lower Rio Grande Valley,
although remarkable that 3 of these, miscogaster, montezuma, and aztecus,
often become abundant here at the northern limit of their range.
In this paper, I have summarized monthly and yearly phaenologic
records accumulated since 1973, indicated those plants most often visited by
Zethus, and provided a 1st report on observations of daily phaenology and
activity temperatures commenced in 1977. Periods available for collection
of Valley Zethus have included, in each year since 1973, 30 days in December
and January, 1 week in March, 2 weeks in May, 2 weeks in June, 1 week in
August, and 1 week in September, as well as 1 week each in April and No-
vember. Between 19-27 November 1977, I sampled Zethus every hour from
0900-1700 and separately labeled each hour's catch. From 18-31 December
1977 this technique was supplemented with maximum-minimum thermom-
eters to record daily temperature ranges and small, portable thermometers

Contribution No. 411, Bureau of Entomology, Division of Plant Industry, Florida De-
partment of Agriculture and Consumer Services, Gainesville, Fla. 32602.
2 Hymenoptera: Eumenidae.
I Research Associate, Florida State Collection of Arthropods, Florida Department of Agri-
culture and Consumer Services, Gainesville.

Vol. 61, No. 3, 1978

The Florida Entomologist

to obtain hourly readings of both sun and shade temperatures at each col-
lecting site.

KEY TO UNITED STATES Zethus
1. Third gastric sternite with a conspicuous translucent apical
lamella or flange that is truncate abruptly on each side before
attaining lateral margin of sternite ......................................... ...... 2
1'. Third gastric sternite with apical flange weakly to conspicu-
ously differentiated but not abbreviated toward lateral mar-
gin of sternite ......................... ........................................... .. ............ 3
2. Hind margin of 2nd gastric tergite evenly convex, hardly
membranous; 1 well developed mid-tibial spur; interocellar
area without tubercles (Texas to El Salvador) .......................
........................ ................... ................ ............. 4. Z aztecus S assure
2'. Hind margin of 2nd gastric tergite divided into 3 sections by
translucent lateral lobes; 2 mid-tibial spurs; interocellar
area with broad, more or less polished tubercles separated by
a narrow line of punctures (Arizona to El Salvador) ....................
....................... .... .... ............ ..................... .. Z guerreroi Z avattari
3. Female clypeus mat with very minute regular longitudinal
striae, as well as numerous large punctures; male flagellum
rolled toward apex; mid-tibia with 1 well developed apical
spur; gaster mostly red (Florida) .............................. Z. slossonae Fox
3'. Female clypeus shining with strong punctures and/or coarse
striations but without minute wrinkling; male flagellum
hooked toward apex; mid-tibia with 1 or 2 apical spurs; gaster
mostly black or in occasional specimens of miscogaster
la r g e ly re d ....................................................... ...... ........ .... ............. 4
4. Stem of 2nd gastric tergite definitely longer than that of 1st;
notaulus weakly impressed or absent (Arizona and Texas to
Argentina)................................... .................... 1. Z. m iscogaster Saussure
4'. Stem of 2nd gastric tergite definitely shorter than that of 1st;
notaulus distinct over at least apical 0.5 of mesoscutum.................... 5
5. Apical propodeal lamella produced into a rounded lobe
above valvula; 2 well developed mid-tibial spurs; petiole
slender (Texas to Colombia) ...................... 2. Z. montezuma Saussure
5'. Apical propodeal lamella scarcely distinguishable from rest
of submarginal carina; only 1 well developed mid-tibial spur;
petiole moderately to very stout..................... .............. .......... 6
6. Height of petiole markedly less than 1/2 the distance from
insertion of flexor muscle to apex; 1st hind tarsomere of
male narrow (Texas to PanamA) .................... 3. Z. otomitus Saussure
6'. Height of petiole equal to 1/2-3/5 the distance from insertion
of flexor muscle to apex; 1st hind tarsomere of male flattened
beneath and conspicuously broadened (eastern United States)
.................... ..... ..... ..... ...... ... ........................... Z sp in ip es S a y
1. Zethus (Z.) miscogaster Saussure (Fig. 1)
MATERIAL EXAMINED. 81 females, 104 males: Bentsen Park, Botanical
Garden, Santa Ana National Wildlife Refuge.
MONTHLY PHAENOLOGY. 7 females and 3 males in January, 1 male in

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Valley Zethus

Fig. 1. Zethus miscogaster. Triangles indicate geographic distribution
in Middle and North America. Fig. 2. Zethus montezuma. Triangles indicate
geographic distribution in Middle and North America.
March, 1 male in May, 2 females and 1 male in June, 10 females and 24
males in August, 25 females and 21 males in September, 24 females and 35
males in November, and 10 females and 11 males in December.
ANNUAL PHAENOLOGY. 2 females and 3 males in 1973, 2 males in 1974,
4 females and 4 males in 1975, 20 females and 16 males in 1976, and 57
females and 77 males in 1977.
TEMPERATURE AND DAILY ACTIVITY PERIOD. From 20-22 November
1977, when maxima and minima in the shade were about 18-28C and skies
ranged from clear to partly cloudy, 1 miscogaster was taken between 0900-
1000, 3 from 1000-1100, 2 from 1100-1200, 3 from 1200-1300, 6 from 1300-
1400, 6 from 1400-1500, 5 from 1500-1600, and 1 from 1600-1700.
Sampling from 22-27 December, when better temperature records were
kept, yielded the following data.
22 December. Minimum -0.60C, maximum in shade 230C; sky clear;
2 males from 1400-1500 (23C shade, 370C sun).
24 December. Minimum 12.90C, maximum in shade 27.50C; partly
cloudy; 1 male from 1100-1200 (220C shade, 290C sun), 2 males from 1300-
1400 (26.50C shade, 390C sun), 2 males from 1400-1500 (27.50C shade, 32C
sun), 1 female and 2 males from 1500-1600 (220C shade, 29"C sun).
25 December. Minimum 12.90C, maximum in shade 25.50C; partly
cloudy; 1 female from 1400-1500 (25.50C shade, 28.50C sun).
27 December. Minimum 100C, maximum in shade 220C; partly cloudy;
1 female from 1200-1300 (220C shade, 33C, sun).
The above data from late fall and early winter 1977 show that misco-
gaster may be active from 0900-1700 and at a shade temperature range of
22-27.50C and at a sun range of 28.5-390C. It flies most abundantly during
early afternoon, when temperatures are highest, and normally seeks herbs,
shrubs, or trees in direct sun.
2. Zethus (Z.) montezuma Saussure (Fig. 2)

The Florida Entomologist

MATERIAL EXAMINED. 71 females, 25 males: Bentsen Park, Botanical
Garden, Santa Ana National Wildlife Refuge.
MONTHLY PHAENOLOGY. 34 females and 7 males in January, 3 females
and 3 males in March, 1 male in April, 2 males in August, 3 females in
September, 11 females and 2 males in November, and 20 females and 10
males in December.
YEARLY PHAENOLOGY. 3 females and 1 male in 1973, 15 females and 6
males in 1974, 28 females and 7 males in 1975, 6 females and 5 males in
1976, and 19 females and 6 males in 1977.
TEMPERATURE AND DAILY ACTIVITY PERIOD. From 21-24 November
1977, when shade minima and maxima were about 10-28C and skies ranged
from clear to partly cloudy, 1 montezuma was netted between 1000 and
1100, 2 from 1100-1200, 2 from 1200-1300, 4 from 1300-1400, and 2 from
1500-1600.
During the December 1977 period of hourly collection and temperature
recording, this species appeared only on the 24th, a warm, partly cloudy
day (minimum 12.90C, maximum in shade 27.5C), when 1 specimen was
registered from 1100-1200 (220C shade, 29C sun), 1 from 1300-1400 (26.50C
shade, 390C sun), and 1 from 1400-1500 (27.50C shade, 320C sun).
During November and December, montezuma may fly from 1000-1600
and at shade temperatures of 22-27.5C and in a sun range of 29-39C. Most
specimens are captured in bright sun and in early afternoon when tem-
peratures are hottest.

3. Zethus (Z) otomitus Saussure (Fig. 3)
MATERIAL EXAMINED. 2 females: Bentsen Park, 1 female, 30-VIII-77;
Botanical Garden, 1 female, 27-VIII-8-IX-73.
FIELD NOTES. My Valley otomitus were netted from flowers of Bac-
charis in bright sun at the edge of dense woods. I have also collected many
in northeast M6xico, mostly in deep, humid forest as they flew around the
dead trunks and logs in which they were nesting. Calmbacher (1977:135-7)
has described the nest of this species.
DISTRIBUTION. Bohart and Stange (1965:118) cite otomitus from no
farther north in M6xico than Nayarit and Veracruz, so the Valley records
provide a noteworthy range extension. I have also found it common in
late May and early June every year from 1974-77 in the densely forested
Cola de Caballo ravine near Monterrey, M6xico, and only 240 km west
and a little south of the Valley.

4. Zethus (Zethoides) aztecus Saussure (Fig. 4)
MATERIAL EXAMINED. 25 females and 32 males: Bentsen Park, Bo-
tanical Garden.
MONTHLY PHAENOLOGY. 1 female and 1 male in January, 1 female in
May, 3 males in June, 2 females and 9 males in August, 13 females and
14 males in September, 4 females and 3 males in November, and 4 females
and 2 males in December.
YEARLY PHAENOLOGY. 7 females and 3 males in 1973, 1 male in 1974,
3 females and 4 males in 1975, 8 females and 21 males in 1976, and 7
females and 3 males in 1977.
TEMPERATURE AND DAILY ACTIVITY PERIOD. On 21 November 1977, I
collected 3 aztecus, of which 1 was taken between 1000 and 1100 and the

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Valley Zethus

Fig. 3. Zethus otomitus. Circles indicate geographic distribution in
Middle and North America. Zethus spinipes. Squares indicate geographic
distribution in Texas. Fig. 4. Zethus aztecus. Rhombs indicate geographic
distribution in Middle and North America.

other 2 from 1300-1400. The day was clear with a shade temperature range
of 18-28C.
Only 1 aztecus was netted during the December sampling period and this
on the 31st (minimum 16.70C, maximum in shade 25.50C) and between
1500 and 1600, when the shade temperature was 250C and the sun reading
270C.
In fall and winter, aztecus thus may be active from 1000-1600, ap-
parently when shade temperatures have reached at least 250C. It almost
always flies in bright sun and becomes most abundant during the warmest
part of the day.

CONCLUSIONS
ZOOGEOGRAPHY. Zethus is a huge genus centered in tropical South
American forests and thorn scrub. It also has a substantial endemic Middle
American radiation plus some species in warmer parts of the Old World.
Only 7 Zethus reach the United States, and 1 ranges north of the Mexican
border area or south Florida. These wasps are well sclerotized, tolerant of
bright sun and high temperatures, and seem able to fly long distances.
Available information suggests that most species also can use a wide va-
riety of caterpillars as prey. Low winter, temperatures consequently may be
the major factor which keeps them from invading more successfully the
deserts of the southwestern United States or the Pine-Oak woods of the
northeast Gulf strip.
Valley Zethus belong to several of the major zoogeographic patterns
displayed by Neotropic biota. Z. miscogaster ranges from Arizona and
south Texas to Argentina and pertains to a group of 6 species, in which 4
are strictly South American and 2 extend over much of South and Middle

The Florida Entomologist

America. Such ichneumonids as Coccygomimus caeruleus Brull, C. sumi-
chrasti Cresson, and Eiphosoma dentator Fabricius, the sphecids Sphex
servillei Lepeletier and Isodontia fuscipennis Fabricius, the vespid Brachy-
gastra lecheguana Latreille, and the papilionid butterflies Papilio anchi-
siades Esper and P. astyalus Godart provide other examples of the same
wide-ranging, endemically South American biogeographic category. Z.
montezuma has been found from Texas to Colombia but pertains to a species
group that ranges south to Paraguay, although centered in Middle America.
The ichneumonids Coccygomimuspunicipes Cresson (Texas to north Chile),
Conopyge spp. (Texas to Brasil) and Eurydacus spp. (M6xico to coastal
Peri) furnish other examples of this northern South American and Middle
American tropical pattern. In contrast, Z. aztecus reaches only El Salva-
dor and forms a monotypic, predominately Middle American group but one
with close relatives in South America. Ichneumonids such as Cryptanura
compact Cresson, C. lamentaria Cameron, Lymeon leucosoma Cameron,
and Polycyrtidea limits Cushman, and the sphecid Trachypus mexicanus
Saussure follow a similar pattern. Finally, Z. otomitus, which ranges from
Texas to PanamA, has a close relative in Colombia, while the other member
of its group, Z. spinipes, occurs in the eastern United States from Massa-
chusetts to Kansas and into Texas as far south as Victoria. Although oto-
mitus and spinipes come within 350 km of each other in east Texas, they do
not intergrade, so their geographic isolation may have been lengthy. This
now mainly Middle and South American species group must have ranged
far north into the eastern United States during warmer Tertiary times, so
that the ancestor of spinipes survived in some Austroriparian refugium
during Pleistocene glacial maxima while those populations that were to
become otomitus retreated into M6xico. Both remained forest adapted,
and thus during interglacial expansion have not reestablished contact
across the south Texas thorn scrub. Remnants of the eastern North American
Neotropic biota are detectable among many groups of organisms. Forty
Neotropic genera of Ichneumonidae, many with species restricted to the
eastern United States, attain at least Maryland (Porter 1977:77-78). Among
the Heteroptera of Connecticut, 33% of the genera show Neotropic affini-
ties (Slater, 1974:153). The southeastern United States even has a few
Neotropic reptiles, such as Rhadinaea flavilata Cope, Tantilla coronata
Baird and Girard, and Anolis carolinensis Voigt.
PHAENOLOGY: as detailed under each species, Valley Zethus follow a uni-
form daily phaenologic regimen. They begin to fly when the shade tempera-
ture is 22C and the sun temperature 27C and during the cooler months
peak in early to mid afternoon and rarely appear before 1000 or after 1600.
All 4 species are thermophilic, but miscogaster, montezuma, and aztecus
easily survive light frosts, as shown by their continued presence in December
and January. I do not have quantitative data on their daily flight periods
in summer, but they seem most abundant from 0900-1200 with less activity
after 1300, when shade temperatures attain 34-35C.
Valley Zethus show substantial differences in monthly phaenology, as
summarized in Table 1.
Z. miscogaster peaks between August and December with maximum
abundance in November. The November figure is significant because I col-
lected in the Valley during that month only in 1977; whereas the data
for May, June, August, September and December derive from 1973-77 and

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Valley Zethus

TABLE 1. MONTHLY PHAENOLOGY OF
RIo GRANDE VALLEY.

Species

Zethus SPECIES FROM THE LOWER

Jan. Mar. Apr. May June Aug. Sept. Nov. Dec.

Z. miscogaster 10 1 1 3 34 56 59 21
Z. montezuma 41 6 1 2 3 13 30
Z. otomitus 2 -
Z.aztecus 2 1 3 11 27 7 6

Total specimens/ 53 7 1 2 6 49 86 79 57
month
Total specimens
collected: 340

those for January and March from 1974-77. Z. aztecus, always less common
than the preceding, has a comparable cycle but reaches greatest abundance
in September, and is relatively rare by November. Z. otomitus has been
collected in the Valley only in August but is common near Monterrey,
Mexico in late May and early June. Z. montezuma is strongly internal,
with 74% of the specimens taken in December and January. Thus, although
all 4 Valley Zethus may coincide at certain times of the year, each species
has a different peak. When we know more about prey preferences and nesting
habits, we may find that this temporal displacement helps reduce compe-
tition for resources such as prey or nesting sites, particularly since all
species frequent the same habitats. However, Z. montezuma, whose optimum
flight period is most out of phase with its congeners, also differs from them
in being smaller (average length to apex of tergite 2 about 11-13 mm in
miscogaster and aztecus and 9-11 mm in montezuma), and this size dif-
ference alone should eliminate spatial and trophic conflicts.
Valley Zethus also fluctuate in numbers from year to year, as docu-
mented in Table 2.
Yearly population density fluctuations of the type noted above are
normal for most Valley insects. Probably they result from the region's
unstable climate characterized in winter by occasional killing frosts and at
any season by droughts which often last 2-3 months. Under these conditions,
few species can be equally abundant from year to year, and some of those

TABLE 2. ANNUAL PHAENOLOGY OF Zethus SPECIES FROM THE LOWER
RIo GRANDE VALLEY.

Species 1973 ,1974 1975 1976 1977

Z. miscogaster 5 2 8 36 134
Z. montezuma 4 21 35 11 25
Z. otomitus 1 1
Z. aztecus 10 1 7 29 10

Total specimens/yr. 20 24 50 76 170

The Florida Entomologist

with strong dispersal powers may disappear from the Valley, only to be
renewed in favorable periods from the more diverse and stable communities
that occupy the lush eastern slopes of the Sierra Madre Oriental in nearby
M6xico.

HABITAT PREFERENCES AND PLANT ASSOCIATIONS: Valley Zethus occur
in all environments from open fields with scattered shrubs to dense gallery
woods but almost always are found in direct sun. The species visit a variety
of plants, where they feed on nectar of flowers, honeydew, or seek twigs
suitable for nesting. All species seem to have about the same range of habi-
tat and plant preferences, some of which are discussed in the following
list of plants with which I have found Valley Zethus most consistently
associated.
Acacia farnesiana L. (Leguminosae). Occasional Zethus, particularly
miscogaster, fly around this shrub at almost any season. The flowers,
which appear mostly in spring, attract a few montezuma but are not reg-
ularly visited by other Zethus species.
Acacia greggii A. Gray (Leguminosae). The flowers of this shrub, which
appear sporadically in winter and last through spring into early summer,
consistently attract miscogaster, montezuma, and aztecus.
Prosopis juliflora DC. (Leguminosae). In spring and early summer the
flowers of this shrub are visited by an occasional Z. montezuma but are not
commonly frequented by any Zethus.
Serjania sp. (Sapindaceae). I have swept a few Z. montezuma from this
ubiquitous vine.
Condalia obovata Hook (Rhamnaceae). During November-January
some specimens of this shrub become very attractive to Z. miscogaster,
montezuma, and aztecus. At this season, C. obovata often is the best source
of Zethus but during the rest of the year is not especially productive.
Bumelia sp. (Sapotaceae). Like C. obovata, this shrub may be visited
by many Zethus during late fall and winter.
Labiatae (unidentified species). This small, white-flowered, scarcely
aromatic mint carpets the forest floor in many areas of the Bentsen Park
and Valley Botanical Garden. During fall and winter specimens in bright
sun may attract numerous Z. miscogaster and montezuma.
Aster spinosus L. (Compositae). Numerous Z. miscogaster were netted
from the flowers of this tall, green-stemmed herb during November 1977.
Aster sp. (Compositae). This brownish-stemmed species also attracted
many miscogaster in November 1977.
Baccharis spp. (Compositae). At least 2 species of these shrubs grow in
the Valley. When they are in bloom, from late August to November,
they attract multitudinous Zethus and other aculeates. This is the only
plant visited by all 4 Valley Zethus and the one which consistently has
been most productive.
All Valley Zethus show about the same floral associations, which vary
seasonally according to which species are blooming or covered with honey-
dew. Shrubs and small trees are the favored plants, and I have collected
Zethus most often in the 2-4 m stratum of Valley woodlands, although
during late fall and winter I have often taken miscogaster and montezuma
from small labiates only 150-300 mm above the ground.

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Valley Zethus

COLLECTIONS
Material covered in this study has been deposited in the Florida State
Collection of Arthropods (Division of Plant Industry, P.O. Box 1269,
Gainesville, Fla. 32602) and in the author's private collection at 301 N.
39th Street, McAllen, Texas 78501.

ACKNOWLEDGMENTS
My Valley fieldwork has been supported in 1976-1977 by United States
National Science Foundation Grant DEB-75-22426 and during 1973-1975 by
grants from the Committee for Research and Exploration of the National
Geographic Society. Collecting permits for Bentsen Park were issued by the
Texas Parks and Wildlife Department (current permit number 1-78), and
Mrs. Vivian Thacker and Mr. A. R. Baker of McAllen, Texas, have facili-
tated fieldwork in the Valley Botanical Garden. Mr. Anthony F. Cerbone
of Fordham University ably assisted me during some of the fieldwork
involved in this project.

LITERATURE CITED
BOHART, R. M., AND L. A. STANGE. 1965. A revision of the genus Zethus
Fabricius in the western hemisphere. Univ. California Pub. Ent.
40:1-208.
CALMBACHER, C. 1977. The nest of Zethus otomitus. Fla. Ent. 60:135-7.
PORTER, C. 1975a. A new Floridian Polycyrtidea with comments on zoo-
geography of Florida Mesostenini. Fla. Ent. 58:247-55.
PORTER, C. 1975b. New records for Zethus from Texas. Fla. Ent. 58:303-6.
PORTER, C. 1977. Ecology, zoogeography, and taxonomy of the Lower
Rio Grande Valley mesostenines. Psyche 84:28-91.
SLATER, J. A. 1974. A preliminary analysis of the derivation of the Heterop-
tera fauna of the northeastern United States with special reference to
the fauna of Connecticut. Connecticut Ent. Soc., 25th Anniversary
Memoirs:145-213.

FIREFLY FLICKER FUSION FREQUENCY.-(Note). Flicker fu-
sion for an adult female of Photuris versicolor (complex) (Coleoptera:
Lampyridae) was found to occur at 41.6 and 45.4 flickers per second in 2
successive trials on the same insect. The stimulus had an absolute energy
level of 0.602 microwatts per square centimeter and a peak intensity at 550
nanometers. The insect had been allowed to dark-adapt for 45 minutes be-
fore testing. Equipment and technique were those used by H. R. Agee (1971,
Ann. Ent. Soc. Amer. 64:942). The test animal, captured as a larva, closed
11 days prior to the study. It has been deposited with Dr. J. E. Lloyd, De-
partment of Entomology and Nematology, University of Florida. I thank
Drs. H. R. Agee (U.S.D.A.), J. E. Lloyd (Univ. Fl.), and J. C. Davis
(U.S.D.A.) for their assistance and use of facilities.-R. M. Merkhofer, De-
partment of Botany, University of Florida, Gainesville 32611.

The Florida Entomologist

BOOK REVIEW

ARTIFICIAL DIETS FOR INSECTS, MITES, AND SPIDERS. Pritam Singh. 1977.
Plenum Publ., New York. xi+594 p., $75. Pritam Singh is among the fore-
most experts in insect rearing and the information contained in his book is
absolutely essential for anyone specializing in this field or directly in-
volved with an organized insectary. Earlier compendia published in co-
operation with H. L. House and W. W. Batsch, and Singh's "Bibliography of
Artificial Diets for Insects and Mites," naturally led to the production of
this single comprehensive volume. The book is impeccably accurate,
thorough, and timely; it ranks with the classical works of A. Peterson
(1934), J. G. Needham et al. (1937), C. T. Brues (1946), and C. N. Smith et al.
(1966). It is considerably more up-to-date than its only contemporary, "In-
sektenzucht" by Winiger (1974), and already has become the standard en-
tomological reference on insect rearing.
This book is totally unpretentious, from its dark brown cover to its
camera-ready format. Actually, it may be considered a large manual. It
begins with a brief historical discussion, literature review, and narrative
glossary (ie., artificial vs. synthetic, holidic vs. meridic, axenic vs. synxenic,
etc.). Singh then follows with the first published account of interactions
among physical factors associated with artificial diets. This explanation in-
cludes diet preparation, nutritional constituents, water content, texture,
and token stimuli. The terse introduction ends with sections on microbial
contamination, the potential role of symbionts, and a scheme for evaluat-
ing diets. Individual citations are arranged alphabetically by taxon and
are listed on the next 467 pages (1961 entries for 754 species-Lepidoptera,
258; Coleoptera, 204; and Diptera, 138; etc.). Each citation consists of an
abstract with sections for diet composition and preparation, rearing pro-
cedures, and typical insect development. Corollaries are indicated for re-
lated references. The volume concludes with an extensive and extremely
useful 74 page set of author and species indices.
Insect rearing has "arrived" as a scientific discipline and, in this respect,
Singh's monograph is a milestone. He listed Bogdonov (1908) as the first
reference to an artificial diet for insects, but only 154 citations are pre-1950.
This means that at least 1807 diets have been developed during the past 25
years. Unfortunately, his literature review was completed just before the
concepts of quality control in insect mass-rearing (Boller and Chambers,
1977) and facilities for insect production (Leppla and Ashley, 1978) were
published. His efforts also preceded the symposium, "Characterization and
Evaluation of Insect Colonies," presented at the XV International Con-
gress of Entomology. However, the only major criticism is the terribly in-
flated price of this relatively austere book. Even though personal owner-
ship has been all but precluded, Singh is to be congratulated for providing
us with access to this splendid document.-N. C. Leppla; USDA; Insect At-
tractants, Behavior and Basic Biology Laboratory; Gainesville, FL, 32601.

Vol. 61, No. 3, 1978

The Florida Entomologist

ECOLOGICAL NOTES ON
LOWER RIO GRANDE VALLEY SPHECINI'P

CHARLES C. PORTER3

Department of Biological Sciences
Fordham University, Bronx, N.Y. 10458

ABSTRACT
Notes on phaenology, distribution, and flower preferences are given for
the 10 species of Sphex, Isodontia, and Prionyx collected by the author
since 1973 in Hidalgo County, Texas. Isodontia fuscipennis (Fabricius) is
recorded as new for the United States, and the first completely documented
United States records of Sphex servillei Lepeletier are adduced.

Since 1973 I have been surveying Hymenoptera in the Lower Rio Grande
Valley of Texas. This work has produced studies of the ichneumonid genus
Thyreodon (Porter 1976) and the eumenid genus Zethus (Porter 1975), as
well as a monograph of the ichneumonid tribe Mesostenini (Porter 1977).
I now add observations on the sphecid tribe Sphecini, whose taxonomy has
been elucidated by Bohart and Menke (1963, 1976) but whose ecology is
as yet poorly documented.
Sphecines are large wasps, strong of flight, and resistant to dessication.
They frequent open areas in bright sun, abound in thorn scrub and deserts,
and, unlike many other Hymenoptera, reach maximum abundance during
the hottest months of the year. Sphecines stock their nests with Orthoptera,
and each genus normally chooses a specific prey family. For example, of
the south Texas genera, Sphex provisions subterranean nests with Tet-
tigoniidae, Prionyx is also fossorial but hunts Acrididae, and Isodontia
stores oecanthine gryllids in hollow plant stems.
Although at the northern limit of the Neotropics, the Lower Rio Grande
Valley has a biota surprisingly similar to that of other subtropical semi-
arid habitats in Middle and South America. Furthermore, it is not far from
the warm-temperate deserts of western North America nor from the humid
Austroriparian Pine-Oak forests of the northeast Gulf strip. The Valley thus
has a zoogeographically complex sphecine fauna of both Sonoran and Neo-
tropic derivation, and its Neotropic element comprises forms of South,
Middle and southeastern North American affinities.
My principal collecting localities in the Valley have been the Bentsen
Rio Grande Valley State Park near Mission and the Valley Botanical
Garden at McAllen. These are designated "Bentsen Park" and "Botanical
Garden" in the accounts of each species given below.

Contribution No. 410, Bureau of Entomology, Division of Plant Industry, Florida Depart-
ment of Agriculture and Consumer Services, Gainesville, Florida 32602.
2 Hymenoptera: Sphecidae.
3 Research Associate, Florida State Collection of Arthropods, Florida Department of
Agriculture and Consumer Services, Gainesville.

Vol. 61, No. 3, 1978

The Florida Entomologist

1. Sphex (S.) ashmeadi (Fernald)
MATERIAL EXAMINED. 4 females, 2 males: BENTSEN PARK, 1 female,
5-VI-73; 1 male, 25-VIII-77; 3 females, 1 male, 31-VIII-77.
FIELD NOTES. Collected in bright sun from flowers of Cissus incisa
Nuttall.
PHAENOLOGY. 1 female in June, 3 females and 2 males in August;
taken only in 1973 and 1977.
2. Sphex (S.) dorsalis Lepeletier
MATERIAL EXAMINED. 7 females, 32 males: BENTSEN PARK, 1 male,
29-VIII-77; 2 males, 31-VIII-77; BOTANICAL GARDEN, 1 male, 21-V-77; 1
female, 1-VI-76; 1 male, 3-VI-76; 5 males, 11-VI-77; 1 female, 4 males,
12-VI-77; 2 males, 2-IX-76; 1 male, 3-IX-77; 1 male, 4-IX-75; 5 males,
4-IX-76; 1 male, 5-IX-76; 1 female, 5 males, 11-IX-76; 1 male, 5-IX-76;
1 female, 5 males, 11-IX-76; 4 females, 2 males, 12-IX-76; 1 male, 27-VIII
to 8-IX-73.
FIELD NOTES. Collected on Cissus incisa, Melilotus alba Desrousseaux,
Pluchea camphorata Linnaeus, and Serjania sp. Twenty specimens (5 fe-
males, 15 males) were taken during September 1976 from flowers of Pluchea
camphorata L. in a low and often flooded part of the Valley Botanical
Garden.
PHAENOLOGY. 1 male in May, 2 females and 10 males in June, 3 males
in August, and 5 females and 17 males in September; 2 males in 1973,
absent in 1974, 1 male in 1975, 6 females and 16 males in 1976, and 1 female
and 14 males in 1977.
Willink (1951:170-1) reported dorsalis as active in northern Argentina
from November to May.

3. Sphex (S.) habenus Say
MATERIAL EXAMINED. 2 females, 8 males: BENTSEN PARK, 1 male,
31-VIII-77; BOTANICAL GARDEN, 1 male, 12-VI-77; 2 females, 3 males,
11-IX-77; 3 males, 12-IX-76.
FIELD NOTES. Collected on flowers of Cissus incisa and occasionally
of Pluchea camphorata, in tall grass at the edge of Celtis woods, and on
Serjania vines in bright sun.
PHAENOLOGY. 1 male in June, 1 male in August, 2 females and 6 males
in September; absent from 1973-1975, 3 males in 1976, 2 females and 5
males in 1977.
4. Sphex (S.) servillei Lepeletier
MATERIAL EXAMINED. 2 males: BENTSEN PARK, 30-VIII-77; 8-IX-76.
FIELD NOTES. Collected on flowers of Baccharis sp. and of Cissus in-
cisa.
PHAENOLOGY. In south Texas appears limited to late summer; no
records for 1973-1975.
Willink (1951:138) indicated that servillei flies from January to April
in northern Argentina.
TAXONOMY. Bohart and Menke omitted servillei from their 1963 re-
vision of the Nearctic Sphecini but cited it, without more specific data, for
"south Texas" in their 1976 checklist (p. 116).
The following diagnosis will separate servillei from other Sphex (S.)

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Sphecini 171

occurring in the Lower Rio Grande Valley: body and legs opaque black
with rather inconspicuous areas of silvery white pubescence on part of
front, most of clypeus, pronotal collar dorsally, apicad on pronotal
lobe, narrowly on apical half of lateral margin of mesoscutum, on part of
postscutellum, more or less of propodeal apex, mesopleuron behind and
below pronotal lobe and again above mid-coxal base, and elongately on
metapleuron from base of hind coxa dorsad along stigmatal groove about
halfway to spiracle; male wings moderately darkened apicad but otherwise
more or less hyaline; female with yellowish or amber-tinged wings; male
7th sternite with a dense tuft of setae on each side and apically emarginate
but without a rounded median projection; male 8th sternite without a
median spine, its apical margin a little acutely rounded.
5. Sphex (S.) texanus Cresson
MATERIAL EXAMINED. 2 females, 7 males: BENTSEN PARK, 2 males, 30-
VIII-77; 1 male, 31-VIII-77; BOTANICAL GARDEN, 1 male, 21-V-77; 1 female,
5-VI-76; 1 female, 1 male, 11-IX-76; 2 males, 12-IX-76.
FIELD NOTES. Collected from flowers of Cissus incisa and from Ser-
jania vines in bright sun.
PHAENOLOGY. 1 male in May, 1 female in June, 3 males in August,
1 female and 3 males in September; absent in 1973-1975, 2 females and 3
males in 1976, 4 males in 1977.
6. Sphex (Fernaldina) lucae (Saussure)
MATERIAL EXAMINED. 3 females, 10 males: BOTANICAL GARDEN, 1 male,
17-24-III-74; 1 female, 20-III-76; 1 female, 16-30-V-74; 1 male, 28-V-75;
1 male, 29-V-75; 1 male, 1-VI-76; 1 female, 1 male, 2-VI-76; 1 male, 4-VI-
76; 2 males, 5-VI-75; 1 male, 9-VI-73; 1 male, 8-IX-74.
FIELD NOTES. Collected mainly on flowers of Melilotus alba. S. lucae
is more sluggish and easily captured than the Valley species of Sphex (S).
PHAENOLOGY. 1 female and 1 male in March, 1 female and 2 males in
May, 1 female and 6 males in June, and 1 male in September; 1 male
in 1973, 2 males and 1 female in 1974, 4 males in 1975, 2 females and 3
males in 1976, absent in 1977.
7. Isodontia fuscipennis (Fabricius)
MATERIAL EXAMINED. 1 female: BENTSEN PARK, 1-13-VI-73.
TAXONOMY. Bohart and Menke described and keyed I. fuscipennis in
their revision of the Nearctic Sphecini (1963:133, 135-7) but cited no records
from the United States.
PHAENOLOGY. Willink (1951:82) reported that this species flies in north-
ern Argentina from December to May.
8. Isodontia elegans (Smith)
MATERIAL EXAMINED. 1 female: BOTANICAL GARDEN, 5-VI-75.
FIELD NOTES. Collected while flying in tall grass at the edge of Celtis
woods.
9. Isodontia mexicana (Saussure)
MATERIAL EXAMINED. 10 females, 22 males: BENTSEN PARK, 2 males,
30-VIII-77; BOTANICAL GARDEN, 1 female, 17-24-III-74; 4 females, 4 males,
16-30-V-74; 2 males, 21-V-77; 1 female, 5 males, 22-V-77; 2 males, 1-VI-76;

The Florida Entomologist

1 male, 2-VI-76; 1 male, 3-VI-76; 1 female, 6-VI-76; 1 male, 11-VI-77;
1 female, 12-VI-77; 1 female, 18-VI-77; 1 female, 22-VIII-77; 1 male, 3-
IX-77; 1 male, 4-IX-77; 1 male, 11-1-76.
FIELD NOTES. Usually found in grassy areas and often on flowers of
Melilotus alba.
PHAENOLOGY. 1 female in March, 5 females and 11 males in May, 3
females and 5 males in June, 1 female and 3 males in August, and 3 males
in September; absent in 1973, 5 females and 4 males in 1974, absent in
1975, 1 female and 6 males in 1976, and 4 females and 12 males in 1977.
10. Prionyx parkeri Bohart and Menke
MATERIAL EXAMINED. 14 females, 32 males: BENTSEN PARK, 1 female,
3 males, 1-13-VI-73; BOTANICAL GARDEN, 1 male, 16-III-74; 3 males, 22-
III-74; 2 males, 17-24-III-74; 1 male, 20-III-76; 5 females, 12 males, 16-
30-V-74; 1 female, 2 males, 25-V-75; 2 males, 28-V-75; 1 female, 29-V-75;
5 males, 1-13-VI-73; 1 female, 5-VI-75; 1 female, 2 males, 7-VI-75; 2 fe-
males, 1-VI-76; 2 females, 2-VI-76; 1 female, 6-VI-76; 1 female, 12-VI-76;
1 male, 1-IX-75.
FIELD NOTES. Collected in tall weeds and overgrown fields, sometimes
on flowers of small herbs, and occasionally on Pluchea camphorata.
PHAENOLOGY. 7 males in March, 7 females and 16 males in May, 7 fe-
males and 11 males in June, and 1 male in September; 8 females in 1973,
5 females and 12 males in 1974, 4 females and 6 males in 1975, 5 females
and 2 males in 1976, and absent in 1977.

CONCLUSIONS
ZOOGEOGRAPHY. The sphecine genera represented in the Valley have es-
sentially cosmopolitan ranges. Sphex is most diverse in the Neotropics
with large radiations also in warmer parts of the Old World but with
fewer species in temperate regions. Isodontia is best developed in the Neo-
tropic and Oriental Realms but has a few species in the temperate Northern
Hemisphere and 2 in Australia. The more xerophile Prionyx occurs almost
everywhere but is best represented in southwest Asia and the Mediterra-
nean zone and has an important radiation in the semiarid Andean and
Chaco regions of South America.
At the specific level, Valley sphecines show either Neotropic or Sonoran
affinities. The Neotropic element includes S. servillei, S. habenus and
S. dorsalis, I. fuscipennis, and P. parkeri; while the Sonoran element is
represented by S. ashmeadi, S. texanus, S. lucae, I. mexicana, and I. elegans.
S. servillei and I. fuscipennis, which range from the Valley to Argentina,
appear to be warm-adapted, originally South American species which have
been prevented by diminishing temperatures from spreading far north into
the United States along the Gulf Arc ,during the present interglacial period.
However, I exornata of the southeastern states is intimately related to
fuscipennis. This suggests that a common ancestor of the fuscipennis-ornata
pair may have occupied the whole Gulf region during warmer Tertiary times.
Later, this ancestor was fragmented by Pleistocene glaciation into eastern
(Florida) and western (Middle America) isolates. The isolates have never
reestablished contact because exornata has adapted to the humid Austrori-
parian Pine-Oak forest and so does not enter the semiarid thorn scrub that

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Sphecini 173

begins southwest of Houston, Texas, while fuscipennis, although hygrically
more versatile, requires subtropical temperatures with at most light and
occasional winter frost.
Similar to the above mentioned, but more cold tolerant, is S. dorsalis,
which extends from Maryland and California to Argentina. S. dorsalis has
not been cited previously from north of Georgia on the Atlantic seaboard,
but I collected it regularly during late summer in 1967-72 at Hudson
near Cambridge, Maryland, on the Delmarva Peninsula. Here it often
visited flowers of Asclepias. Such immense intercontinental distributions
are not uncommon among euryhygric, vagile Neotropic insects. For ex-
ample, the mesostenine ichneumonids Acerastes pertinax Cresson and
Pachysomoides stupidus Cresson or such diurnal Lepidoptera as Battus
polydamas Linnaeus, Eurema nicippe Cramer, Phoebis philea Linnaeus,
P. sennae Linnaeus, Danaus gilippus Cramer, Heliconius charitonius Lin-
naeus, and Agraulis vanilla Linnaeus have ranges more or less comparable
to that of S. dorsalis. As pointed out in my review of Valley Mesostenini
(Porter 1977:78), few Neotropic insects are "tropical" in the sense of re-
quiring frost-free winters, so that those which can cross the arid zones of
northern M6xico and the southwestern United States often attain 35 to
400 N. Lat.
Although the Neotropic biota originated in South America, it also now
flourishes almost intact in most of Middle America. The huge Middle
American Neotropic element obviously is the product of multiple in-
vasions, which occurred at various times throughout the Tertiary and
warmer periods of the Pleistocene, so that it consists not only of species
like S. servillei and I. fuscipennis, which extend uniformly to southern
South America, but also of many species that have evolved locally from
South American ancestors and in response to the orogeny and severe cli-
matic changes that affected the region during later Tertiary and Pleistocene
times. Prionyx parkeri seems to fit this latter category. It belongs to the
Thomae group, which is centered in the semiarid Andean and Chaco regions
of South America (6 endemic species and 2 more that reach northward into
the United States), with P. parkeri in most of the United States and Mexico
south to the Isthmus of Tehuantepec, 1 species generally distributed over
the United States and northern M6xico, 1 well represented in the western
United States with scattered records from the east, and another confined to
the west. The Thomae group in North America consequently shows a So-
noran facies-i.e., species centered in drier parts of the west. However, true
Sonoran elements are most diverse in the North and Middle American
semiarid zones, with secondary invasions of South America (e.g., the meso-
stenine ichneumonid genus Compsocryptus, Porter 1977:80), while the
Thomae group predominates in semiarid southern South America and has a
subsidiary radiation in M6xico and western North America. We may thus
regard this group as part of a genuinely xerophile Neotropic biota, in con-
trast to the main hygrophile Neotropic element that consists of such forest
insects as Ichneumonidae, social Vespidae, etc. This assemblage is analo-
gous to the Sonoran but arose in South America, probably, as envisioned
by Solbrig (1976:36), starting in the early Tertiary and "at middle latitudes,
particularly in the western part of the continent" where a "flora adapted
to a seasonally dry climate" has long existed. Xerothermic episodes later
in the Tertiary and in the Pleistocene allowed northward movement of

The Florida Entomologist

this element, just as they permitted some Sonoran genera to reach South
America. The nyssonine sphecid tribe Bembicini also largely corresponds
to this distributional pattern, as do such bee genera as Caupolicana, Dia-
dasia, and Centris, and even a few ichneumonids, such as the Planosae
group of Trachysphyrus (Porter 1977:81).
Finally, S. habenus appears to be a northeast North American Neo-
tropic element which became distinct from originally South American an-
cestors while isolated during Pleistocene glacial maxima in Florida or the
Gulf states and which in the present interglacial has moved southwest
again as far as Mexico. In this regard, it resembles many ichneumonids,
such as the mesostenines Diapetimorpha macula Cameron, D. introita
Cresson, D. acadia Cushman, and Lymeon orbus Say (Porter 1977:77).
As pointed out by Halffter (1976:28-29), the more or less xeric Mexican
and western North American Sonoran fauna seems composed of some
elements that evolved from ancient South American ancestors which moved
north in the late Cretaceous and early Tertiary and of others stemming from
equally old predecessors that crossed the Bering land bridge from Asia in
those same climatically benign times. These taxa now are centered in the
Sonoran geographic area but many have invaded Central or South America
while others have reached the southeastern United States or even the West
Indies. Some of these wider-ranging Sonoran forms have remained xero-
philic and show disjunct distributions in peripheral areas now under rela-
tively humid climates (e.g., the ichneumonid genus Compsocryptus and the
snake genus Pituophis), while others have evolved a few widely distributed
forms adapted to conditions wetter and/or cooler than those prevailing
in the place of origin (e.g., the mesostenine genus Joppidium and the snake
genus Crotalus). All the above distributional types are found among the
Valley Sonoran sphecines. Sphex ashmeadi and S. texanus, both confined
to the western United States and northern M6xico, seem, like the other
North American Sphex (S.), to be of ultimate South American derivation.
On the other hand, Sphex (Fernaldina) lucae, of general distribution in the
western United States and M6xico as far south as Guadalajara and with
some isolated records in the southeastern United States also, has only 1
close relative, S. (F.) melanocnemis of the eastern Mediterranean area.
This disjunct pattern may suggest early Tertiary dispersal around the
Northern Hemisphere with subsequent extinction in many areas and sur-
vival by adaptation to aridity. At any rate, the subgenus Fernaldina is
absent from South America. Finally, I. elegans (western United States and
northern M6xico) and L mexicana (eastern United States to Wyoming,
Arizona, and all of Middle America) belongs to a group also represented
by 1 species that is restricted to the eastern United States and 1 found only
in Cuba (Bohart and Menke 1976:120). None of this assemblage reaches
South America, and its closest relatives are in the Oriental and eastern
Palaearctic areas. These facts may indicate a trans-Bering movement in
the early Tertiary with subsequent mid to late Tertiary differentiation in
the arid and mountainous parts of western North America, southward move-
ment into Middle America during Pleistocene glacial maxima, and various
northward and eastward pulses in glacial minima.
The Lower Rio Grande Valley sphecine fauna treated in this study thus
is made up of 5 Neotropic and 5 Sonoran species belonging to cosmopolitan
genera. The species all are widely distributed, and none is endemic to the

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Sphecini 175

Valley. Four of them (S. ashmeadi, S. texanus, S. lucae, and L elegans) are
shared principally with the rest of the western United States and northern
M6xico, 1 (S. habenus) with the southeastern United States and northern
Mexico, 2 (I. mexicana and P. parkeri) are widely distributed in the United
States and Middle America, 2 (S. servillei and I fuscipennis) extend from
the Valley to Argentina, and 1 (S. dorsalis) from Maryland to Argentina.
PHAENOLOGY. I have collected in the Valley both with hand nets and
Malaise traps but have taken sphecines there only by hand. Periods avail-
able for hand collecting each year since 1973 have included approximately
30 days in December and January, 1 week in March, 2 weeks in May, 2 weeks
in June, 1 week in August, and 1 week in September, as well as 1 week each
in April (1975) and November (1977). Sphecines were collected only in
March, May, June, August, and September, and these records are summarized
in Table 1.
Valley sphecines thus fly mostly from late spring to early fall, with
1-15 June being the optimum period sampled (9 species, 47 specimens).
Their numbers and diversity build up steadily from March to June but have
diminished by the last part of August only to increase dramatically again
during the first week of September. On the other hand, intensive collecting
in December and January has failed to obtain any Sphecini.
This monthly phaenology is approximated by numerous Valley Hymen-
optera and in particular by almost all the Sphecidae. However, most other
Valley warm season Hymenoptera show an even more pronounced summer
hiatus than the Sphecini. On the other hand, the only sphecids abundant in
winter are several species of Liris, although occasional specimens of Cer-
ceris spp., Ochleroptera bipunctata Say, Trachypus mexicanus Cameron,
and Trypoxylon spp. appear as late as December or January and a large
sphecid fauna still may be active toward the end of November.

TABLE 1. MONTHLY PHAENOLOGY OF LOWER RIo GRANDE VALLEY
SPHECINI.

Mar. May June Aug. Sept. Tot.
Sp.
1. S. ashmeadi 1 5 6
2. S. dorsalis 1 12 3 22 38
3. S. habenus 1 1 8 10
4. S. servillei 1 1 2
5. S. texanus 1 1 3 4 9
6. S.lucae 2 3 7 1 13
7. fuscipennis 1 1
8. elegans 1 1
9. mexicana 1 16 8 4 3 32
10. P. parkeri 7 23 15 1 46
Total species/
month 3 5 9 6 7
Total specimens/
month 10 44 47 15 40
Total specimens collected: 156

The Florida Entomologist

In contrast, many Valley Hymenoptera remain active throughout the
benign winters, and some of the parasitic groups attain maximum abundance
and diversity in December and January. For example, mesostenines peak in
December-January but become rare in summer (Porter 1977:82-83), and
the same is true for most other Ichneumonidae and for certain Chalcidoidea,
particularly the genus Spilochalcis. The winter fauna also includes nu-
merous braconids; the scoliid Campsomeris tolteca Saussure; Zethus,
Pachodynerus, Hypalastoroides, Leptochilus, Stenodynerus, and Eumenes
among the Eumenidae; the vespids Mischocyttarus, Brachygastra, and
Polistes; such pompilids as Ageniella, Anoplius, Auplopus, Calicurgus, Cry-
ptocheilus, Minagenia, and Priocnessus; and diverse bees, including
Hylaeus, Agapostemon, Augochlora, Temnosoma, Halictus, Dialictus,
Megachile, Coelioxys, Exomalopsis, Xylocopa, and Bombus.
We have no evidence as to why some Valley Hymenoptera are adult
principally in winter and others mainly from March to November or as to
why so many of the warm season taxa decline in summer. Probably, in most
cases the reason is trophic rather than climatic. Large orthopterans hunted
by sphecines are most abundant during the warm season, whereas the holo-
metabolous immatures utilized by parasitoids, scoliids, eumenids, and
many vespids tend to be most available in winter, and the spiders caught
by Pompilidae on the flowers visited by bees are present in the Valley at
any season. On the other hand, extreme summer heat, droughts, and oc-
casional winter frosts also could be important population controls, but
experimental evidence is needed for their evaluation. Certainly, climatic
factors often must act in complex, subtle ways, and phaenology cannot be
predicted from zoogeographic affinities. For example, the mainly South and
Middle American S. servillei and I fuscipennis fly, as might be expected,
during summer, but many other "tropical" Hymenoptera of similar distri-
bution (ichneumonids such as Coccygomimus caeruleus Brull6, Lymeon
leucosoma Cameron, Bicristella texana Porter, Cryptanura lamentaria
Cameron, C. vallis Porter, and Conopyge conica Brull6; the eumenid
Zethus montezuma Saussure; and the halictid Temnosoma sp.) in the
Valley peak or occur exclusively in winter.
Within their observed activity period, some Valley sphecines are most
common toward late summer and others in spring. The 5 species of Sphex
(S.) all peak in August or September, and none occurs earlier than May.
On the other hand, S. (F.) lucae, I mexicana, and P. parkeri peak in May or
June, emerge in March, and have become rare by September, while I. fusci-
pennis and I. elegans are known only for June. There seems to be no ob-
vious relationship between this temporal differentiation and any trophic
or habitat factors by which it would reduce competition among the species
affected.
Although collecting was done each year at approximately the same
times and in the same localities, numbers and diversity of species varied
strikingly from year to year. These data are summarized in Table 2.
It is noteworthy that none of the species was collected every year, that
I. fuscipennis and I. elegans appeared only once, and that 1976 and 1977
yielded by far the greatest numbers and diversity. As explained elsewhere
(Porter 1977:85), the Valley enjoys a subtropical climate that is violently
altered at intervals by protracted droughts, short periods of torrential
rain and flooding, and occasional killing frosts. These density independent

Vol. 61, No. 3, 1978

Porter: Lower Rio Grande Sphecini 177

TABLE 2. ANNUAL PHAENOLOGY OF LOWER Rfo GRANDE VALLEY SPHE-
CINI.
1973 1974 1975 1976 1977

1. S. ashmeadi 1 5
2. S. dorsalis 2 -1 22 15
3. S. habenus -- 3 7
4. S. servillei 1 1
5. S. texanus 5 4
6. S. lucae 1 3 4 5
7. L fuscipennis 1 -
8. L elegans 1 -
9. I. mexicana 9 7 16
10. P.parkeri 8 17 10 7
Total species/year 5 3 4 7 6
Total specimens/year 13 28 16 48 48

factors probably explain most of the fluctuations recorded in Table 2.
For example, the productive summer of 1976 was unusually wet and fol-
lowed a rather wet winter. In 1977, almost equally productive, the winter
was one of the wettest on record and led into a particularly verdant spring
followed by almost 3 months of drought and then a short period of abun-
dant rain in September, when vegetation and insects revived dramatically.
Most sphecines thus seem favored by more humid periods in the Valley's
erratic pluvial spectrum. However, S. (F.) lucae and P. parkeri were absent
for the first time in 1977, perhaps because these more xerophile species
suffered disproportionately in the damp, cool winter of 1976-1977.
FLOWER PREFERENCES. Adult sphecines regularly drink nectar of
flowers. Available data, although incomplete, suggest considerable differ-
entiation in flower preference among Valley species.
Cissus incisa (Vitaceae) is the best plant for Sphex (Sphex) and attracts
all 5 Valley species, including fair numbers of ashmeadi, dorsalis, habenus,
and texanus.
Melilotus alba (Leguminosae) is visited by a few S. dorsalis and by many
S. lucae and I. mexicana.
Pluchea camphorata (Compositae) yields large quantities of S. dorsalis,
an occasional S. habenus, and some P. parkeri.
Baccharis spp. (Compositae), although especially productive for most
anthophilous Hymenoptera, have produced only S. servillei.

COLLECTIONS
SMaterial used in this study has been divided between the Florida State
Collection of Arthropods (Division of Plant Industry, P.O. Box 1269,
Gainesville, Florida 32602) and the author's private collection (301 N. 39th
St., McAllen, Texas 78501).

ACKNOWLEDGMENTS
My Valley fieldwork has been supported in 1976-1977 by United States
National Science Foundation Grant DEB-75-22426 and during 1973-1975 by

The Florida Entomologist

grants from the Committee for Research and Exploration of the National
Geographic Society. The Texas Parks and Wildlife Department has issued
permits for insect collecting in the Bentsen Rio Grande Valley State Park
(current permit number 1-78), and Mrs. Vivian Thacker and Mr. A. R. Baker
of McAllen, Texas, have facilitated fieldwork in the Valley Botanical
Garden. Mr. Anthony F. Cerbone of Fordham University has assisted me
during some of the fieldwork involved in this study.

LITERATURE CITED
BOHART, R. M., AND A. S. MENKE. 1963. A reclassification of the Sphecinae.
Univ. Cal. Pub. Ent. 30:91-182.
BOHART, R. M., AND A. S. MENKE. 1976. Sphecid wasps of the world. Uni-
versity of California Press, Berkeley, Los Angeles, London. p. 1-695.
HALFFTER, G. 1976. Distribuci6n de los insects en la zona de transici6n
mexicana. Relaciones con la entomofauna de Norteam6rica. Folia
Entomologica Mexicana 35:1-64.
PORTER, C. 1975. New records for Zethus from Texas. Fla. Ent. 58:303-6.
PORTER, C. 1976. New records for Thyreodon from south Texas. Psyche
83:304-9.
PORTER, C. 1977. Ecology, zoogeography, and taxonomy of the Lower Rio
Grande Valley mesostenines. Psyche 84:28-91.
SOLBRIG, 0. 1976. The origin and floristic affinities of the South American
temperate desert and semidesert regions, p. 7-49 In Goodall, D. (ed.),
Evolution of desert biota. University of Texas Press, Austin.
WILLINK, A. 1951. Las species argentinas y chilenas de Chlorionini. Acta
Zool. Lilloana 11:53-225.

Vol. 61, No. 3, 1978

The Florida Entomologist

FORMICA INTEGRA 2. FEEDING, TROPHALLAXIS,
AND INTERSPECIFIC CONFRONTATION BEHAVIOR'2

R. C. WILKINSON3, A. P. BHATKAR', W. J. KLOFT4,
W. H. WHITCOMB3, AND E. S. KLOFT4
ABSTRACT
In laboratory screening tests, Formica integra Nylander attended
honeydew-producing insects such as the aphid, Neosymydobius albasiphus
Davis, on swamp chestnut oak and a soft scale, Toumeyella parvicornis
(Cockerell), on slash pine seedlings. Forest insect pest species consumed by
F. integra included Neodiprion excitans Rohwer and N. lecontei (Fitch),
Ips calligraphus (Germar) and grandicollis (Eichh.), Reticulitermes flav-
ipes (Kollar), Rhyacionia frustrana (Comstock), and Tetralopha robustella
Zeller. Trophallaxis between F. integra workers taken from the same or
widely-separated nests in the field was demonstrated by use of a radioiso-
tope. When 3 potentially competitive ant species were paired with F. integra
in laboratory confrontation tests and their mortality was compared with
that of F. integra by a calculated confrontation index, mortality was twice
as much for Crematogaster atkinsoni Wheeler, one-half as much for Solen-
opsis invicta Buren, and only one-fifth as much for Camponotus abdomi-
nalis floridanus (Buckley).

Habitat, nest construction, polygyny and biometry of a red wood ant,
Formica integra Nylander 1856, from west-central Georgia were described
in a preceding paper by Kloft et al. (1973). We conducted further studies
during 1972-74 on ants from the same area to estimate the potential use-
fulness of F. integra as a predator of pest species such as pine sawflies.

FEEDING BEHAVIOR
Feeding habits of certain species of red wood ants (especially Formica
polyctena Foerster in Germany) have been reported in detail in the Euro-
pean literature. English abstracts of papers between 1930 and 1961 were
given by Cotti (1963); Way (1963) reviewed mutualism between ants and
honeydew-producing Homoptera. Adlung (1966) and Finnegan (1971) dis-
cussed ant feeding habits in regard to effective use of these predators as
biological control agents. Red wood ants attend certain species of psyllids,
coccids, and aphids (Kloft et al. 1965) to obtain honeydew, which is re-
ported to contain carbohydrates and proteins in amounts sufficient to main-
tain colonies during periods when insect prey is scarce. Red wood ants are
reported to prey most effectively on hairless caterpillars and sawfly
larvae which defoliate hardwood or coniferous trees, but many other kinds
of arthropods are also carried to ant nests for consumption. Feeding and
foraging activities of F. polyctena are mostly diurnal; however, Horstmann

Florida Agricultural Experiment Station Journal Series No. 755.
Hymenoptera: Formicidae.
SDepartment of Entomology and Nematology, University of Florida, Gainesville, 32611.
SInstitute for Applied Zoology, University of Bonn, West Germany.

Vol. 61, No. 3, 1978

The Florida Entomologist

(1970) found that such activities were primarily temperature-dependent, and
29% of total honeydew collected plus 14% of total prey were collected
at night.
In Manitoba, Canada, Bradley and Hinks (1968) found that Formica
spp. (especially Formica obscuripes Forel) attended Cinara spp. aphids
[especially Cinara gracilis (Wilson)] infesting jack pine, Pinus banksiana
Lamb. In the study area, F. obscuripes was seen carrying geometrid, tor-
tricid, and noctuid larvae, adult psyllids and beetles, and parts of grass-
hoppers, flies, and spiders along the trails to their nests. Many of the larvae
were defoliators of jack pine and most were still alive while being carried.
Two successful attempts have been made by Finnegan (1975, 1977b) to intro-
duce Formica lugubris Zetterstedt 1840 from Italy and F. obscuripes from
Manitoba into Quebec, Canada. There he found these ants readily preyed on
larvae of diprionid sawflies such as Neodiprion lecontei (Fitch) and N.
swainei Middleton (Finnegan, personal communication), and on the spruce
budworm, Choristoneura fumiferana (Clemens) (Finnegan 1977a).
We made observations in Georgia and Florida to determine whether
F. integra would attend sucking insects and prey on different species of
insects offered to it. Special attention was paid to predation of the sawflies,
Neodiprion excitans Rohwer and N. lecontei, 2 important defoliators of
pines in Florida (Hetrick 1959, Wilkinson 1969).
METHODS.-Field observations were made at the site in Georgia described by
Kloft et al. (1973). F. integra worker ants from Georgia were transferred
to a laboratory at Gainesville, Florida and established in a 1-m diam by
20-cm deep plastic "nest arena" containing nest material from Georgia on
moistened sand. The nest arena was connected by 2 large plexiglass tubes
to a similar "feeding arena" in which artificial ant diet (Bhatkar and Whit-
comb 1970), 10% honey-water, and water were offered daily. The laboratory
room was maintained at 27 + 20C, 50-80% RH, and light:dark cycle of
14:10 h. Worker activity during dark periods was observed by use of a red
light. Live insect prey, or plants or branches infested with insects, were
placed in the feeding arena to observe feeding-foraging activities of F. in-
tegra. At least 10 specimens of each species were offered as prey in each
trial.
RESULTS.-F. integra workers attended and obtained honeydew from
Neosymydobius spp. aphids feeding on 2 closely-related species of chestnut
oaks growing in Georgia and Florida (Table 1.). Honeydew was also ob-
tained from a soft scale, Toumeyella parvicornis (Cockerell), which is com-
monly found on young pines in Florida. Naked, unprotected insects (in-
cluding many pest species) were preyed upon in most cases, but webbing
readily protected larvae of the pine webworm, Tetralopha robustella
Zeller. F. integra workers did not open sawfly cocoons, but carried off
live sawflies and sawfly parasites. Both N. excitans and N. lecontei sawfly
larvae were constantly harassed by F. integra workers, were unable to feed,
and eventually fell to the soil in' the feeding arena. Fallen larvae
were immediately seized and carried into the nest arena. Ant feeding-
foraging activity was noticeably greater during light periods, but attendance
of the soft pine scale and oak aphids as well as harassment and predation
of feeding sawfly larvae continued at a reduced rate during dark periods.
DIscussIoN.-Foraging and predation activities of F. integra under labora-
tory conditions were very similar to those reported for European and

Vol. 61, No. 3, 1978

Wilkinson et al.: Formica integra

Canadian species of red wood ants under laboratory and field conditions.
Attendance of Neosymydobius sp. aphids by F. integra in the field in Georgia
(southern edge of F. integra range) and the readiness of F. integra to utilize
a related Neosymydobius sp. farther south in Florida are desirable traits in
relation to potential use of F. integra as a biological control agent (cf.
Finnegan 1971, 1975). This relationship might be exploited in Florida by
introducing F. integra into loblolly pine, Pinus taeda L., sawtimber
stands where swamp chestnut oak, Quercus michauxii Nutt., is often present
and is commonly infested with Neosymydobius spp. aphids. Outbreaks of
N. excitans sawflies commonly occur in such sawtimber stands in Florida
(Wilkinson, unpubl.).

TROPHALLAXIS
A typical behavior of red wood ants is the distribution of liquid food
from 1 individual to another by regurgitation (trophallaxis), so that all
individuals in a nest share food gathered by foraging workers. This could be
an important factor in successful establishment and maintenance of col-
onies in new areas (Finnegan 1971). Food distribution in certain European
species has been studied with radioactive tracers (Goesswald and Kloft
1956, 1958, 1960, 1963; Kneitz 1963; Lange 1958, 1960). Similar experiments
were conducted to determine whether trophallaxis occurs in F. integra.
METHOD.-In all 3 experiments, radioactive sugar-water (20%, labelled as
2"P orthophosphate, specific activity 0.3 mCi/ml, was offered to 1 worker
which fed to the point of satiation. After careful external decontamination
by washing in buffered phosphate "chaser" solution (Kloft 1977), the sati-
ated ant (donor) was placed with a group of 54-58 workers which had been
removed from a laboratory nest and starved for 4 h (acceptors). Three single
experiments involved (1) a donor from nest "A" and 58 acceptors from nest
"A"; test ended after 4.5 h, (2) a donor from nest "A" and 58 acceptors from
nest "A"; test ended after 20 h, and (3) a donor from nest "A" and 54 ac-
ceptors from nest "B"; test ended after 4.5 h (Fig. 1). Experiments were ter-
minated after 4.5 or 20 h by killing all ants with chloroform. The dead
workers were individually checked for radioactivity measured in counts per
minute (cpm), which was considered to be proportional to an individual's
participation in the chain transfer of liquid food initiated by a single donor.
Workers from laboratory nests "A" and "B" were originally obtained in
Georgia from nests located about 200 m apart in the field.
RESULTS.-The cpm values for individual workers are grouped in logarith-
mic classes in Fig. 1. Both donors and acceptors in experiments 1 and 2 came
from laboratory nest "A" and only the observation times differed. A fairly
normal distribution was obtained in experiment 1 after 4.5 h, even though
8 out of the 58 acceptors had still received practically no food (0-10 cpm).
The peak of radioactivity in experiment 2 shifted to higher values after 20 h
indicating that all individuals had participated in trophallaxis. Transfer
of radioactivity in experiment 3 from a donor in nest "A" to nearly all
of the 54 potential receptors from nest "B" corresponded with the results of
experiment 1.
DIscussION.-Trophallaxis between F. integra workers taken from the same
or widely separated nests in the field was demonstrated under laboratory
conditions; results were similar to those obtained by laboratory and field

182 The Florida Entomologist Vol. 61, No. 3, 1978

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Wilkinson et al.: Formica integra

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The Florida Entomologist

1 11- NEST A --- 5801 NEST A Hs
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0-10 11-100 101-1000 1001-10000 10001-100000
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Fig. 1. Transfer of radioactivity among F. integra workers from 3 single
donor ants fed on "P sugar-water to 54-58 receptor ants per each of 3 rep-
licates after 4.5 or 20 h. (1) Solid symbol: transfer from donor from nest
"A" to acceptors from nest "A" after 4.5 h; (2) Cross-hatch symbol: transfer
from donor from nest "A" to acceptors from nest "A" after 20 h; (3) Hollow
symbol: transfer from donor from nest "A" to acceptors from nest "B" after
4.5 h.
experiments with F. polyctena in Europe (Goesswald and Kloft 1963, Kloft
1967, Lange 1960). Such involvement of large populations in a food chain
should increase survival and efficiency of red wood ants as predators. We
expect that this situation applies to F. integra, but it remains to be proved
under field conditions.

INTERSPECIFIC CONFRONTATION BEHAVIOR
Interspecific aggression is a basic drive in ants and may vary from soli-
tary aggression in more primitive forms such as Myrmeciinae, to aggressive
recruitment and mass attack in more advanced forms such as Formicinae
and Myrmicinae (Robertson 1971). Aggression can be pronounced when
species compete for the same nesting sites and forage for the same food at
the same time. Our confrontation tests were conducted to determine whether
F. integra (native to Georgia) might compete successfully if introduced into
territory inhabited by 3 species: (1) Crematogaster atkinsoni Wheeler
(native to Florida), (2) the red imported fire ant, Solenopsis invicta Buren
(introduced and widely established in Florida), and (3) the Florida car-
penter ant, Camponotus abdominalis floridanus (Buckley) (native to

Vol. 61, No. 3, 1978

Wilkinson et al.: Formica integra

Florida). F. integra might be expected to compete with these 3 species,
since they attend some of the same species of honeydew-producing insects
[e.g., C. atkinsoni also attends Neosymydobius albasiphus Davis aphids on
oak, while both C. atkinsoni and C. a. floridanus attend T. parvicornis
soft scale on pine (see feeding behavior)]. All 4 species forage during
daylight. F. integra occurs in terrestrial nests in semi-open woods, while
C. atkinsoni is found in arboreal nests and S. invicta in terrestrial nests
in the open. Pronounced aggression might be expected between F. integra
and C. a. floridanus, which both nest in rotting logs and stumps in semi-
open woods.
METHOD.-Simple confrontation tests (Bhatkar et al. 1972) were conducted
in a lighted room at a mean 280C and 65% RH. Confrontation units con-
sisted of 2 plastic petri dishes 5.5 cm diam x 1.5 cm high, lined with moist
filter paper and interconnected with 4 mm diam tubing. F. integra was
placed in 1 dish and an alien species in the other. Preliminary tests with
variable numbers of workers per dish indicated that a conclusive outcome
within 24 h could be obtained by pairing workers as follows: 100 F. in-
tegra vs. 200 C. atkinsoni; 100 F. integra vs. 100 S. invicta; 300 F. integra
vs. 100 C. a. floridanus. Final mortality tests (Table 2) involved 10 rep-
licates each of these 3 pairings and deaths were recorded after 24 h. Con-
frontation indices for F. integra were calculated as follows: average mor-
tality for alien species divided by average mortality for F. integra. Auxil-
iary behavioral tests consisted of observing reactions of F. integra workers
to introduction of a probe and to manual jarring of dishes. The reactions
of 20 workers confronted in dishes by an equal number of alien workers
were also observed for 20 min.

TABLE 2. MORTALITY AND CONFRONTATION INDEX FOR Formica integra
WORKERS AFTER 24-H CONFRONTATIONS WITH Crematogaster
atkinsoni, Solenopsis invicta, AND Camponotus abdominalis
floridanus WORKERS.

Workers Mortality
Paired per
species dish* Min.-Max. Avg. Index**

C. atkinsoni 200 138-188 167.3
F. integra 100 75-93 83.4
S. invictat 100 27-48 37.5 0.52
F. integra 100 45-97 71.5
C. floridanus t 100 42-54 49.5 0.20
F. integra 300 247-256 251.2

*n= 10 pairs of dishes per test. No mortality occurred in 5 control dishes of each species.
**Confrontation index (for F. integra)=average mortality of alien species + average mor-
tality of F. integra.
fWe (W. J. and E. S. Kloft) observed interspecific food transfer from 1 C. floridanus
worker to 1 attacking S. invicta worker during 1977, using the tracer methods of Bhatkar and
Kloft (1977). Such "appeasement" behavior might reduce mortality in natural conflicts.

The Florida Entomologist

RESULTS.-F. integra workers reacted to introduction of a probe and man-
ually-induced vibrations in dishes as follows: (1) opened their jaws, (2)
stopped quickly, (3) held their antennae in "U"-shape, parallel to each
other and upwards, (4) moved head and forelegs towards source of distur-
bance, (5) curled abdomen underneath legs and raised forelegs slightly up-
wards, (6) simultaneously moved head and directed abdomen towards
source of disturbance, (7) simultaneously raised forelegs and curled ab-
domen farther so that the terminal acidopore was directed toward point of
disturbance, and (8) sprayed through acidopore.
The principle response of F. integra workers confronted by equal num-
bers of C. atkinsoni, S. invicta, or C. a. floridanus workers also consisted
of spraying through the acidopore. F. integra workers repeatedly responded
as follows: (1) raised their heads and antennae upwards and moved towards
alien workers, (2) opened jaws and jerked bodies, and (3) sprayed alien
workers either with or without seizure of alien workers' appendages or
petiole. Both C. atkinsoni and S. invicta workers often showed crippled
movement and died after 4 or more sprays, but few C. a. floridanus workers
were killed by spray alone.
The principal responses to confrontation with F. integra workers varied
according to species: C. atkinsoni seized legs and smeared venom from the
tip of the abdomen; S. invicta seized appendages, smeared venom, stung
opponents, and dismembered them; C. a. floridanus seized opponents' legs,
petioles, or heads and sprayed.
In comparison with mortality inflicted upon F. integra workers, about
twice as many C. atkinsoni, one-half as many S. invicta, and only one-
fifth as many C. a. floridanus workers were killed by F. integra (Table 2).
DIscussION.-Results suggested that F. integra workers could overcome
C. atkinsoni workers, but not S. invicta or especially C. a. floridanus
workers. In Georgia, F. integra had established numerous nests concentrated
in an area of about 0.1 ha (Kloft et al. 1973). Large, well-established
colonies might ward off S. invicta or C. a. floridanus attacks in Florida.
Finnegan (1975) ensured a more rapid establishment of introduced F. lugu-
bris populations in Quebec by treating competing indigenous Formica and
Camponotus species with insecticide.

ACKNOWLEDGMENTS
The authors thank W. F. Buren, G. W. Dekle, L. A. Hetrick, and A. N.
Tissot for identification of ants, scales, termites, and aphids, respectively,
and R. A. Nickle for graphic illustrations.

LITERATURE CITED
ADLUNG, K. G. 1966. A critical evaluation of the European research on use
of red wood ants (Formica rufa group) for the protection of forests
against harmful insects. Z. angew. Ent. 57:167-89.
BHATKAR, A., AND W. H. WHITCOMB. 1970. Artificial diet for rearing various
species of ants. Fla. Ent. 53:229-32.
BHATKAR, A., W. H. WHITCOMB, W. F. BUREN, P. CALLAHAN, AND T. CAR-
LYSLE. 1972. Confrontation behavior between Lasius neoniger
(Hymenoptera: Formicidae) and the imported fire ant. Environ. Ent.
1:274-9.

Vol. 61, No. 3, 1978

Wilkinson et al.: Formica integra

BHATKAR, A. P., AND W. J. KLOFT. 1977. Evidence, using radioactive phos-
phorus, of interspecific food exchange in ants. Nature 265:140-2.
BRADLEY, G. A., AND J. D. HINKS. 1968. Ants, aphids, and jack pine in
Manitoba. Can. Ent. 100:40-50.
COTTI, G. 1963. Bibliografia ragionata 1930-1961 del Gruppo Formica rufa
in Italiano, Deutch, English. Minist. Agric. Forest. Roma, Coll.
Verde 8, 413 p.
FINNEGAN, R. J. 1971. An appraisal of indigenous ants as limiting agents
of forest pests in Quebec. Can. Ent. 103:1489-93.
FINNEGAN, R. J. 1975. Introduction of a predacious red wood ant, Formica
lugubris (Hymenoptera: Formicidae), from Italy to eastern Canada.
Can. Ent. 107:1271-4.
FINNEGAN, R. J. 1977a. Predation de Choristoneura fumiferana par Formica
lugubris. Rev. bimestrielle de recherches Peches et Environ. Can.
33:1.
FINNEGAN, R. J. 1977b. Establishment of a predacious red wood ant, For-
mica obscuripes (Hymenoptera: Formicidae) from Manitoba to east-
ern Canada. Can. Ent. 109:1145-8.
GOESSWALD, K., AND W. KLOFT. 1956. Untersuchungen ueber die Verteilung
von radioaktiv markiertem Futter im Volk der Kleinen roten Wald-
ameise (Formica rufopratensis minor). Waldhygiene 1:200-2.
GOESSWALD, K., AND W. KLOFT. 1958. Radioaktive Isotope zur Erforschung
des Staatenlebens der Insekten. Umschau 58:743-5.
GOESSWALD, K., AND W. KLOFT. 1960. Untersuchungen mit radioaktiven Iso-
topen an Waldameisen. Entomophaga 5:33-41.
GOESSWALD, K., AND W. KLOFT. 1963. Tracer experiments on food exchange
in ants and termites. Pages 25-42 in IAEA Editorial Staff, eds. Radia-
tion and radioisotopes applied to insects of agricultural importance.
Int. Atomic Energy Agency, Vienna.
HETRICK, L. A. 1959. Ecology of the pine sawfly, Neodiprion excitans
(Rohwer) (Hymenoptera: Diprionidae). Fla. Ent. 42:159-62.
HORSTMANN, K. 1970. Untersuchungen uber der Waldameisen (Formica
polyctena Foerster) im Eichenwald. Oecologia (Berl.) 5:138-57.
KLOFT, W. 1967. Radioedologische Untersuchungen an Formiciden des
Waldes. Wiss. Z. Univ. Dresden 16:582-3.
KLOFT, W. J. 1977. Part V. Pages 141-200 in E. A. Beck and H. L. Cromroy,
eds. Laboratory training manual on the use of isotopes and radiation
in entomology. Int. Atomic Energy Agency, rev. 2nd edit., Vienna.
KLOFT, W., A. MAURIZIO, AND W. KAESER. 1965. Das Waldhonigbuch.
Ehrenwirth Verlag Muenchen. 218 p., Illus.
KLOFT, W., R. C. WILKINSON, W. H. WHITCOMB, AND E. S. KLOFT. 1973.
Formica integra (Hymenoptera: Formicidae) 1. Habitat, Nest Con-
struction, Polygyny, and Biometry. Fla. Ent. 56:67-76.
KNEITZ, G. 1963. Tracerversuche zur Futterverteilung bei Waldameissen.
Symposia Genetica et Biologica Italica XII:38-50.
LANGE, R. 1958. Einfluss der Koenigin auf die Futterverteilung im Ameisen-
staat. Naturwiss. 45:196-7.
LANGE, R. 1960. Ueber die Futterweitergabe zwischen Angehoerigen vers-
chiedener Waldameisenstaaten (Zugleich ein Beitrag zum Problem
des Nestduftes bei den Ameisen). Z. Tierpsychol. 17:389-401.
ROBERTSON, P. L. 1971. Pheromones involved in aggression behavior in the
ant, Myrmecia gulosa. J. Insect Physiol. 17:691-715.
WAY, M. J. 1963. Mutualism between ants and honeydew-producing Homop-
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WILKINSON, R. C. 1969. Control of the red-headed pine sawfly. Sunshine
State Agr. Rep. Nov. 1969. p. 13-5.

The Florida Entomologist

BOOK REVIEW
VIRUS-INSECT RELATIONSHIPS. By K. M. Smith. Longman Inc., New York.
291 pp. Profusely illustrated. $23.50. Written from a biological point of view,
this book provides a wealth of information on viruses that are known to
produce diseases in insects. The book is divided into 2 parts. Part 1, "The
different types of insect viruses", is further subdivided: Part A includes
chapters on the inclusion-type virus diseases, and Part B presents chapters
on the non inclusion-type viruses. The chapters in Part 1 are well organized,
and provide concise descriptions of the virus particles, their polyhedra
(where applicable), the disease caused, host range, and geographic distribu-
tion. Sometimes additional information is given about virus replication,
serology, and transmission. Many electron micrographs show purified virus
particles (usually negatively stained preparations) and thin sections of
virus infected host tissue. Generally, the electron mircographs are excellent
reproductions from papers cited by the author. Diagrammatic representa-
tions showing virus replication and assembly processes are also given for
some of the viruses.
Part 2, "Further aspects of the study of virus-insect relationships", con-
tains 12 chapters that cover a wide range of subjects: routes of infection
and reproduction of insect viruses (Ch. 8), serology of insect viruses (Ch.
9), latent infections (Ch. 11), tissue culture of insect viruses (Ch. 12), and
insects and other arthropods as vectors of plant and animal diseases (Ch. 17,
18). These chapters are quite general, but specific examples from the litera-
ture have been used to consolidate subject matter. Chapters on synthetic
feeding and mass rearing of insects (Ch. 16) and insects and other arthro-
pods as vectors of animal diseases (Ch. 18) are relatively superficial. In view
of the voluminous literature on each of these topics such as cursory review
could have been omitted. The section on staining methods for optical micros-
copy (Ch. 15) could have been greatly enhanced by the addition of brief
descriptions of methodology for study of viruses by electron microscopy.
The book concludes with an appendix on virus diseases of mites, a bibli-
ography containing about 750 references and an index to authors and
subjects.
Knowing Dr. Smith's prominence and reputation as one of the world's
leading insect virologists, I expected more than this volume appears to offer.
However, the book has many strong points. The organization and presenta-
tion of Part 1, and the excellent electron micrographs provide a ready source
of biological information concerning nearly all of the common, and some of
the more unusual insect viruses. Also, the extensive bibliography gives an
excellent entry into the literature on most phases of insect virology. The
book is essentially a review of the literature. In some chapters, the author
has integrated the literature citations into cohesive accounts; other chapters
are not so well integrated.
The book contains at least 1 serious error. While describing the host
range of the iridescent virus affecting Aedes taneniorhychus (p. 107, lines
4-7) the author states "Larvae of Corethrella brageleyi (Diptera: Chaobor-
idae) in Louisiana are voracious predators of first-instar mosquito larvae
and easily pick up the iridescent virus from MIV mosquitoes. The virus was
readily transmissible to both C. brakeleyi and C. appendiculata (Chapman
et al., 1971)." This statement gives the erroneous impression that the irides-
cent virus from Ae. taeniorhynchus and that from C. brakeleyi are the same.
Actually, the paper cited (Chapman et al. 1971. J. Invert. Pathol. 18:284-6)
describes a new iridescent virus from C. brakeleyi. This was cross trans-
mitted to larvae of C. appendiculata, but it COULD NOT be transmitted to
larvae of Ae. taeniorhynchus, Ae. sollicitans, or Psorophora ferox.
Although Virus-Insect Relationships is not a book for virologists inter-
ested in biochemical aspects of insect viruses, it can be a valuable addition
to the library of entomologists and insect pathologists working in biological
control of insects. Some instructors may find Part 1 of the book helpful as
an overview of insect viruses in courses concerning insect pathology.-
Darrell W. Anthony, USDA, SEA, FR; Insects Affecting Man and Animals
Research Lab. Gainesville, FL 32604

Vol. 61, No. 3, 1978

The Florida Entomologist

A 12-VOLT DC POWER PACK FOR ELECTROCUTOR
GRID TRAPS'

E. R. MITCHELL, E. W. HAMILTON, AND S. MASUDA

Insect Attractants, Behavior, and Basic Biology Research Laboratory,
Science and Education Administration, USDA, Gainesville, Fla. 32604

ABSTRACT
A 12-V DC battery-powered unit for electrocutor grids was as effective
in the field as the standard 115-120 V AC powered grid for capturing 6
species of Lepidoptera. The trap operated effectively for an average of 14
nights between battery charges.

The cylindrical electrocutor grid described by Mitchell et al. (1972)
is highly effective in knocking down many different moth species, e.g., the
cabbage looper, Trichoplusia ni (Hubner), soybean looper, Pseudoplusia
includes (Walker), tobacco budworm, Heliothis virescens (F.), corn ear-
worm, H. zea (Boddie), fall armyworm, Spodoptera frugiperda (J. E.
Smith), beet armyworm, S. exigua (Hilbner), and tobacco hornworm,
Manduca sexta (L.). Thus, traps with an electrocutor grid have received
wide acceptance as a research and survey tool, though operation from a
conventional 120 V AC power source does not permit random placement in
remote areas. This problem was solved partially by development of a
portable grid trap operated from a 12 V DC heavy-duty automobile battery
with a commercially available DC to AC inverter (Mitchell et al. 1973).
Although this unit is as effective as the standard AC powered grid, an opera-
tional time of only 28-30 h before the battery must be recharged has limited
its use.
Reported here are results of field tests conducted during the summer of
1977 with the Bodine Model 12BZ power-pack unit (The Bodine Co., 236
Mt. Pleasant Road, Collierville, Tenn. 38017) designed to operate the
electrocutor grid much longer before recharging of the battery becomes
necessary. This unit provides 7000 V DC across the grid rather than the 5000
V AC that the standard grid trap supplies.

METHODS AND MATERIALS
Previous experiences with automobile batteries showed that they were
incapable of withstanding repeated deep discharge-charge cycles when used
as a power source for the electrocutor grid. After 3 or 4 discharge-charge
cycles, an automobile battery would seldom hold an adequate operational
charge. Therefore, we used a Hurricane marine battery (Hurricane Battery
Co., Tampa, Fla.) rated at 70 A in these tests. Marine batteries are designed
specifically for deep discharge and recharge cycles and they are useable
for a much longer time than automobile batteries under these conditions.

SReceived for publication 15 February 1978. This paper reports the results of research only.
Mention of a commercial or proprietary product does not constitute endorsement of that product
by the USDA.

Vol. 61, No. 3, 1978

The Florida Entomologist

A very reliable light-sensing switch device incorporating a photoconductive
cell (TIL 64) was developed to electrically disconnect the battery from
the Bodine power-pack during daylight hours (Fig. 1). The current drain of
this device during the daylight "OFF" time was minimal (ca. 50 pA).

6 VDC RELAY

82 -- 100 mA COIL

12 VDC 2N2484

+12VDC
? TIL64 LOAD

- O -- 4 -------
Fig. 1. Schematic drawing for control circuit activated by a photo-
conductive cell.
The electrocutor traps, one with a grid powered by 115-120-V AC and the
other with a grid powered by the Bodine unit, were compared for their
ability to capture the corn earworm (CEW, Test 1), fall armyworm (FAW,
Test 2), beet armyworm (BAW, Test 3), tobacco hornworm (THW, Test 4),
and the cabbage looper and soybean looper (CL and SBL, Test 5). The
traps in each test were placed next to plantings of corn and tomatoes at
intervals of ca. 20 m and baited with either 4 virgin laboratory-reared
females (CEW, BAW, and THW) or 25 mg synthetic pheromone (FAW,
(Z)-9-dodecen-l-ol acetate; CL and SBL, (Z)-7-dodecen-l-ol acetate) in a
1.25-ml plastic vial. In tests 6 (CEW) and 7 (FAW), introduction of a
switching circuit to rapidly apply and remove power to the Bodine unit
resulted in a pulsating high-voltage output that conserved battery power.
In each test, the traps were checked every 1-2 days, the number of captured
insects was recorded, and the positions of the traps were rotated. Each col-
lection was therefore considered a replicate, and the data were analyzed
by using Student's t-test.
RESULTS AND DISCUSSION
The numbers of moths captured in the grid trap powered by the unmodi-
fied Bodine unit were comparable to the numbers of moths captured in the

Vol. 61, No. 3, 1978

Mitchell et al.: Power Pack for Grid Traps

grid trap powered by standard 120 V AC (Table 1). However, the traps pow-
ered by the modified Bodine unit that switched rapidly on and off captured
significantly fewer corn earworm and fall armyworm moths than the traps
powered by AC.
The unmodified unit was operated for 81 nights (ca. 10 h each), from early
July until the 1st week of October. During this period, the battery's voltage
and specific gravity were checked daily except on weekends. The battery
was changed and recharged when the specific gravity dropped below 1.135
even though the voltage remained relatively high (ca. 11 V). Under these
conditions, trap operation averaged 14 nights between battery charges.
TABLE 1. MEAN NUMBER OF MALE MOTHS CAPTURED PER REPLICATE (+
SE) IN CYLINDRICAL ELECTRIC GRID TRAPS POWERED BY A
STANDARD 115-120 V AC OR A 12 V DC POWER UNIT (BODINE
MODEL 12BZ), ALACHUA, FLA. 1977.*
Mean no. male moths captured in
No. indicated trap
Species Test no. replicates Standard AC Bodine DC

Corn earworm 1 11 49.6+15.2 38.5+ 15.9
Fall armyworm 2 5 186.820.4 155.0 16.8
Beet armyworm 3 24 16.0+ 5.5 24.8 + 7.3
Tobacco hornworm 4 22 4.2 + 0.6 4.1 0.6
Cabbage looper 5 13 3.9+ 0.8 3.5 + 0.9
Soybean looper 5 13 23.8 + 6.8 29.8 5.3
Corn earworm 6 6 22.5+ 3.7 6.0 2.6
Fall armyworm 7 6 173.3 +21.4 71.3 + 10.0

*Differences between results from traps powered by standard AC and those powered by Bodine
DC were significant (Student's t-test, 5% level) only for tests 6 and 7, in which the Bodine
unit was equipped with a pulsing unit to prolong battery life.
Electrocutor grid traps powered by the Bodine unit provide more flexi-
bility than the AC powered unit and are more reliable and less expensive
to operate than the battery-powered unit using a DC to AC inverter (Mit-
chell et al. 1973). It is expected that the battery-powered grid described here
could become a valuable research and survey tool.

ACKNOWLEDGMENT
We gratefully acknowledge the following personnel of this labora-
tory: for assistance in insect rearing-J. Rye and C. Greene; for collecting
insects from the traps-W. Copeland and R. Hines; and for assembling
the photoswitch-F. Lee. We also thank A. Baumhover and other personnel
of the Insect Rearing Unit, Tobacco Research Laboratory, Oxford, N.C., for
supplying some of the bait insects.

LITERATURE CITED
MITCHELL, E. R., J. C. WEBB, A. H. BAUMHOVER, R. W. HINES, J. M.
STANLEY, R. G. ENDRIS, D. A. LINDQUIST, AND S. MASUDA. 1972.
Evaluation of cylindrical electric grids as pheromone traps for
loopers and tobacco hornworms. Environ. Ent. 1:365-8.
MITCHELL, E. R., J. C. WEBB, J. C. BENNER, JR., AND J. M. STANLEY. 1973.
A portable cylindrical electric grid trap. J. Econ. Ent. 5:1232-3.

The Florida Entomologist

AN APPARATUS FOR THE OBSERVATION OF LIVING IM-
MATURE AND SMALL ADULT SPIDERS-(Note). Investigations
of spider development, morphology and physiology sometimes require
that specimens be examined under a microscope and later released alive
and unharmed. This can be accomplished by anesthetizing the spider, but I
believe this has the potential to alter the animal's behavior, physiology,
etc. Such alterations in behavior or physiology are difficult to measure, but
the possibility of their existence must be assumed until data to the contrary
are presented.
The ideal situation is immobilization of the spider without anesthesia
during the period of observation. Seligy (1970, Can. J. Zool. 48:406-7) de-
vised an apparatus for this purpose employing an adjustable piston with a
glass top. The spider was placed in the piston and an adjusting screw
turned, pressing the spider against the glass, immobilizing it without crush-
ing it. Berry, Miller and Harris (1978, Ann. Ent. Soc. Amer. 71:126-8) de-
signed and built a chilling table for immobilizing insects.
I devised a simple apparatus that immobilizes spiders, requires little
material and only a few minutes to fabricate. A 35 x 10 mm tissue culture
dish (Falcon #3001) and some cardboard are required. Use the bottom of the
dish as a template and cut out a circular piece of white paper (to serve as a
background for observations) and tape it in the bottom of the dish. Use the
top of the dish as a template and cut out a circular piece of cardboard. Cut
a circle out of the center of the cardboard circle. Place the cardboard ring
on top of the inverted dish bottom and place the top of the dish over the
ring (Fig. 1) forming the observation chamber.
The depth of the chamber, which should be slightly greater than the
thickness of the specimen, can be adjusted by adding layers of rings of vary-
ing thickness. The mobility of the spider can be further limited by reducing
the diameter of the center of the cardboard rings. The entire apparatus is
easily positioned under a microscope.-John B. Randall, Department of
Entomology and Nematology, University of Florida, Gainesville, 32611.

Lid

SCardboard
ring

Bottom,
inverted

Fig. 1. A cardboard and culture dish apparatus for immobilizing and ob-
serving living spiders.

Vol. 61, No. 3, 1978

The Florida Entomologist

A NEW SPECIES OF CECIDOMYIA INJURIOUS TO
CONES OF SLASH PINE IN FLORIDA'

RAYMOND J. GAGNA

Systematic Entomology Laboratory, IIBIII, Federal Research,
Sci. & Educ. Admin., USDA2

ABSTRACT
A new species of pitch midge, Cecidomyia bisetosa Gagn6, injurious to
cones of slash pine in Florida is described and compared to its closest
congener. Unlike other Cecidomyia, which live in pitch on twigs and
branches of conifers, C. bisetosa attacks the cones producing hypertrophy
of the cone scales and preventing normal seed release from mature cones.

The genus Cecidomyia contains 12 known species, 9 from North America
and 3 from Eurasia, all of which feed on resin of pines and, in Europe,
fir and spruce. The new species described here is the first found to feed on
cones; all others have been reported only from pitch masses on twigs and
branches where they promote breakage and secondary infection. On cones
the damage is more serious because scales become enlarged and deformed
thus preventing release of the seeds when cones mature.
The new species is known so far only from a seed orchard of slash pine,
Pinus elliottii Englemann, in Nassau Co., Florida, but is a potentially
serious threat to seed production in slash pine seed orchards in the southern
United States. It was originally submitted to me for identification by E. P.
Merkel and I. L. Williams, entomologists with the Southeastern Forest
Experiment Station, Olustee, Florida. These entomologists are investigat-
ing the biology of the new species.
C. bisetosa was discovered too late to be included in my revision of
Cecidomyia (Gagnd, in press) but can be referred to the keys that I pre-
pared for that paper. In the key to adults, C. bisetosa will run to couplet
5: sternum X is deeply bilobed, although not as much so as in C. resini-
cola (Osten Sacken), and the aedeagus is parallel-sided. In the key to
larvae, C. bisetosa will run to couplet 6 and differs from resinicola and
the other 2 species in that couplet in its lack of a short, peglike seta on
the terminal abdominal segment.
C. bisetosa is closely related to C. resinicola, a species that is wide-
spread in North America on many different hosts of hard pines including
P. elliottii. The pupal taxonomic characters are alike in both bisetosa and
resinicola, and the male terminalia differ slightly with sternum X of
bisetosa less deeply bilobed and the aedeagus narrower. Larvae of bisetosa
have lost the short peglike seta of the third terminal papilla, and their
hind spiracles are bilaterally symmetrical (compare Figs. 3-4).

SDiptera: Cecidomyiidae.
2 Mailing address: c/o U.S. National Museum, Washington, D.C. 20560.

Vol. 61, No. 3, 1978

The Florida Entomologist

1I ,
II

'-S -.

3 4
Fig. 1-3, Cecidomyia bisetosa Gagn6, n. sp. 1, male terminalia (dorsal);
2, same (lateral); 3, instar III hind spiracle and terminal segments (dorsal).
Fig. 4, Cecidomyia resinicola, instar III hind spiracle and terminal segments
(dorsal).
Cecidomyia bisetosa Gagn6, NEW SPECIES
Adult. Palpus four-segmented. Legs covered with brown scales. Male
terminalia (Figs. 1-2): telomere short; sternum X bilobed, the lobes short;
aedeagus shorter than sternum X, wide, parallel-sided, convex caudally.
Pupa. Antennal horn short with ventral ridge; pronotum with mem-
branous anterior tubercle at each side of center line and 1 long and 1
short seta at its base.
Larva instarr III). Spatula absent. Pleural papillae and lateral pairs
of dorsal papillae of abdomen with short setae not situated on tubercles;
middle pair of dorsal papillae not apparent. Two terminal papillae
present (Fig. 3): 1 with long, tapered seta, one with long, peglike seta.
Terminal spiracle bilaterally symmetrical, with long, caudal prongs.
Holotype, male, ex cones Pinus elliottii, Nassau Co., Florida collected
6-XII-1976, R. Mantie, emerged 2-1-1977, USNM Type No. 75227. Paratypes:

Vol. 61, No. 3, 1978

Gagn6: New Cecidomyia Species

5 males, 4 pupal exuvia, 10 larvae, same data as holotype (USNM);
3 males, 5 females, same host and locality, collected 8-X-1976, I. Williams
(Florida State Collection of Arthropods); 6 larvae, same host and lo-
cality, 1-XII-1977, R. Mantie (USNM).
The specific epithet, bisetosa, refers to the presence of only 2 setae on
each side of the terminal abdominal segment of the larva.

LITERATURE CITED
GAGNt, RAYMOND J. (In press). A systematic analysis of the pine pitch
midges, Cecidomyia spp. (Diptera: Cecidomyiidae). Technical Bulle-
tin. Agr. Res. Serv., USDA.

SUSTAINING ASSOCIATES

Lear Pest Control Co.
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The Florida Entomologist

NEW MEMBERS 1977-78

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Vol. 61, No. 3, 1978