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(ISSN 0015-4040)
FLORIDA ENTOMOLOGIST
(An International Journal for the Americae)
Volume 66, No. 3 September, 1985
TABLE OF CONTENTS
BARANOWSKI, R. M., R. I. SAILER, AND F. W. MEAD-Euschistus
acuminatus, A Pentatomid New to the United States (Hemip-
tera; Pentatomidae) ............................................................................ 287
WmTH, W. W.-The North American Predaceous Midges of the Bezzia
bicolor Group (Diptera: Ceratopogonidae) ....................................- 292
CHILuES, C. C., AND A. G. SELHIM-Reduced Efficacy of Fenbutatin-
oxide in Combination with Petroleum Oil in Controlling the
Citrus Rust Mite Phyllocoptruta oleivora ........................................ 310
ELVIN, M. K., J. L. STIMAC, AND W. H. WHITCOMB-Estimating Rates
of Arthropod Predation on Velvetbean Caterpillar Larvae in
Soybeans ._-_..-.......---------------.. ..--...--..--- .........---.....------- 819
BURK, T.-Behavioral Ecology of Mating in the Caribbean Fruit Fly,
Anastrepha suspense (Loew) (Diptera: Tephritidae) ................ 330
PoRTE, C. C.-Biological Notes on Lower Rio Grande Valley Augo-
chloropsis and Agapostemon (Hymenoptera: Halictidae) ............ 344
POnTER, C. C.-Eiphosoma dentator (Fabricius) in Florida (Hy-
menoptera: Ichneumonidae) -.........................................................-- 353
Scientific Notes
SANTIAGO-BLAY, J. A.-Plumeria rubra: A New Host Plant
Record of the Fall Armyworm (Lepidoptera: Noctuidae)
in Puerto Rico ...................................-....................................... 359
DAY, J. F., AND G. A. CURTIS-Opportunistic Blood-feeding on
Egg-laying Sea Turtles by Salt Marsh Mosquitoes
(Diptera: Culicidae) ........................................................... 359
Frrz, G. N.-A Technique for Separating Mole Crickets from
Soil .....-.--. .....----........................... ........-... --------.......... ... 360
KHALAF, K. T.-Lyriform Organs in the Centipede Scutigera
(Scutigeromorpha: Scutigeridae) ........................................ 362
EHRMAN, L., AND M. WASSERMAN-Firefly Chromosomes
(Photinus, Photuris: Lampyridae, Coleoptera) ---...---....... 363
DIXON, W. N.-Damage of Slash Pine Female Strobili by Repro-
duction W eevils ---................................................................-- .-- 364
Book Review .........----...----.......... ...------- --------------------------------. 366
North American Benthological Society .......................................................- 371
ISI Launches First Data Management System for Reprints ................-.. 372
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This issue mailed November 5, 1983
Baranowski et al.: Euschistus acuminatus 287
EUSCHISTUS ACUMINATUS, A PENTATOMID NEW TO
THE UNITED STATES (HEMIPTERA: PENTATOMIDAE)
R. M. BARANOWSKI, R. I. SAILER AND F. W. MEAD
The University of Florida Agricultural Research and
Education Center, Homestead, FL 33031
University of Florida, Department of Entomology and
Nematology, Gainesville, FL 32611
Florida Department of Agriculture and Consumer Services,
Division of Plant Industry, Gainesville, FL 32611 USA, respectively
ABSTRACT
Euschistus acuminatus Walker is reported for the first time for the
United States. The various stages are described, the 5th instar is figured,
and the biology is discussed.
RESUME
Se registra Euschistus acuminatus Walker por primera vez en los Estados
Unidos. Se described los estadios, se ilustra el quinto instar y se discute la
biologia.
The first Florida specimens of Euschistus acuminatus Walker were
collected by Mr. George Gwinn and Ms. Connie Stieger, Florida Department
of Agriculture and Consumer Services, Division of Plant Industry, from
Cestrum diurnum L. (day jessamine, Solanaceae) at 8301 NW 8th St.,
Miami, FL on 25-VII-1980. These specimens were sent to the Division of
Plant Industry, Taxonomic Section, Gainesville, for identification. Examina-
tion by Drs. F. W. Mead and R. I. Sailer proved them to represent a species
not known from the United States. Specimens were subsequently sent to
Dr. L. H. Rolston, Department of Entomology, Louisiana State University,
Baton Rouge, LA, who identified them as E. acuminatus.
Additional specimens were collected for life history studies by Mr.
E. Pefia, also with the Division of Plant Industry, on 27-X-1980 and by Mr.
George Gwinn on 29-X-1981 at the same location. On the last date, a 4 block
area was surveyed, but E. acuminatus was apparently still confined to the
original site. Cestrum diurnum was also examined in several areas ca.
20-30 miles south of Miami but E. acuminatus was not found. According to
Long and Lakela (1971) C. diurnum has been introduced into Florida from
tropical America. It is found in cultivation throughout south Florida and
has also escaped cultivation.
Euschistus acuminatus can be recognized by the following description.
Head, corium, scutellum and pronotum uniformly yellowish to dark
brown. Corium, scutellum and portion of pronotum posterior to an irregular
transverse line between the humeri with uniform dark punctures; pronotum
anterior to this line and head with punctures concolorous with background
color or only slightly darker. Anterolateral margins of pronotum concave,
denticulate, a black band originating at anterior corner of margin, fading
approximately half way to humeri. Humeral angles strongly produced,
spinose, black, slightly upturned. A prominent black C-shaped and 3 ir-
Florida Entomologist 66 (3)
regular black marks located on either side of the midline on the anterior
portion of the pronotum. Membrane smoky brown, antennae yellowish, legs
yellowish with occasional light brown spots. Venter usually paler than
dorsum. Thoracic pleura typically with 5 black spots on each side.
Adults examined range from 8.8 to 11.2 mm in length.
The prominent black spinose humeral angles and C-shaped marks on the
pronotum readily distinguish acuminatus from all other Florida species of
Euschistus and it will key out in Blatchley (1926) to E. crassus. E.
acuminatus can easily be separated from crassus by the humeral angles
being more strongly produced and by the black marks on the pronotum. The
adult is figured by Barber (1939).
Outside of Florida acuminatus is known to occur in Hispanola (from
where it was originally described), Puerto Rico, (Barber 1939) and Cuba
(Barber and Bruner 1932).
Description of Euschistus acuminatus nymphs
All measurements are in mm.
5TH INSTAR (Lab reared; in alcohol) (N=3), (Fig. 1)
Ground color yellow-brown. Head, pronotum, scutellum and wing pads
punctate and with red to brown dots. Cicatrices on pronotum outlined in
brown. Anterolateral margin of pronotum with small irregular serrations.
Abdominal terga with irregular reddish spots, lateral margin of each tergite
with a U-shaped brown mark; plates surrounding scent gland ostioles with
dark brown irregular spots. Legs yellow-brown, base of setae on femora and
tibiae brown. Antennal segments 1, 2 and distal half of 4 yellow brown with
bases of setae on 1, 2 brown; segment 3 and proximal half of 4 stramineous.
Head length 1.73, width 1.74, interocular space 1.07. Pronotum length 1.39,
width 4.91; wing pad length 2.48; abdomen length 4.45. Labial segments
length I 1.28, II 1.29, III 1.49, IV 1.69. Antennal segments length I 0.60, II
1.16, III 1.19, IV 1.17. Total body length 8.35.
4TH INSTAR (Lab reared, in alcohol) (N=2)
Ground color yellow brown. Head, pro, meso and metanotum punctate;
punctures on disk, scutellar area of mesonotum and all on metanotum except
for lateral ends, surrounded by reddish dots remaining punctures with brown
dots. Pronotal cicatrices outlined in brown, a pair of elongate brown bars
near anterior margin of mesonotum. Abdominal markings similar to 5th
instar. Legs yellow brown, base of setae on femora and tibiae brown; distal
end of tibiae slightly infuscated, tarsi brownish except for mid area of 1st
segment. Antennae yellowish except for brown distal half of segment 4;
base of setae of segments 1, 2 brownish. Head length 1.28, width 1.31,
interocular space 0.92. Pronotum length 0'.81, width 2.96; wing pad length
0.90, abdomen length 2.0. Labial segments length I 0.85, II 0.89, III 0.44, IV
0.44. Antennal segments length I 0.48, II 1.09, III 0.88, IV 0.93. Total body
length 4.78.
3RD INSTAR (Lab reared, in alcohol) (N=1)
Ground color of head, thorax and abdomen cream. Head, thorax with
288
September, 1983
Baranowski et al.: Euschistus acuminatus
Fig. 1. Euschistus acuminatus, 5th instar nymph, dorsal view.
brownish punctures, suture between jugum and tylus brown; a brown
elongate spot mesad of eye margin. Pronotal disc with an elongate brown
ring, on either side of mid line, opened on posterolateral corner. Mesonotum
with 2 elongate bars and metanotum with a single irregular bar in the same
location. Abdominal segments with reddish dots; lateral margin of each
segment with a brown U-shaped mark. Legs brown except for distal half
of femora yellowish. Antennal segments, 1 mostly brown, base of 2 brown,
remaining portion of 2, all of 3 yellowish with base of setae brown, segment
289
Florida Entomologist 66 (3)
4 except for pale base dark brown. Head length 1.0, width 1.06, interocular
space 0.72. Pronotum length 0.50, width 1.88; abdomen length 1.60. Labial
segments length I 0.52, II 0.60, III 0.32, IV 0.38. Antennal segments length
I 0.26, II 0.66, III 0.58, IV 0.70. Total body length 3.60.
2ND INSTAR (Lab reared, in alcohol) (N=1)
Ground color of head, thorax and abdomen whitish. Posterior margin of
head black connecting to an anterior projecting black line mesad of eye
margin extending just anterior to eye margin; also mid line black, may or
may not be connected to black posterior margin. Margins of jugae, suture
between jugum and tylus black. Areas between black lines with numerous
black punctures. Thorax with numerous black punctures, those along pos-
terior margin coalescing; median punctures also connected by irregular black
marking; explanate margins translucent with black markings. Abdomen
with red dots; scent gland plates blackish-brown with whitish mid line and
lateral dots. Legs blackish red except for distal end of femora white. Head
length 0.64, width 0.72, interocular space 0.52. Pronotum length 0.26, width
1.12; length abdomen 0.74. Labial segments length I 0.36, II 0.36, III 0.20, IV
0.26. Antennal segments length I 0.16, II 0.36, III 0.36, IV 0.48. Total body
length 1.80.
1ST INSTAR (Lab reared, in alcohol) (N= 1)
Head light brown with irregular mid line and area posterior to eye
cream colored. Thorax uniformly brown with cream colored mid line.
Abdomen reddish with irregular whitish spots coalescing around brownish
scent gland plates, margin of abdominal segments with brownish triangular
spots. Legs brownish, with apical end of tibiae and tarsi cream. Antennae
burgandy with intersegmental areas red. Head length 0.60, width 0.58,
interocular space 0.38. Pronotum length 0.16, width 0.84; Abdomen length
0.74. Labial segments length I 0.18, II 0.18, III 0.14, IV 0.22. Antennal seg-
ments length I 0.10, II 0.20, III 0.20, IV 0.36. Total body length 1.40.
EGG (Lab reared, in alcohol) (N= 10)
Typically barrel shaped, somewhat more flattened on the anterior end, a
uniform pale green color when first deposited, changing to tan in alcohol;
34-40 slender micropylar projections, 0.08 in length and having the distal
end somewhat thickened, arranged around the pseudoperculum. Chorion
with a triangular reticulated pattern with a fine spine or setae at each angle.
LABORATORY STUDIES
Stock specimens of E. acuminatus were kept in plastic containers 120 mm
in dia, 180 mm high with a screened top. 'Tap water was provided in a vial
with a cotton wick. Plant material was changed every 3-5 days. To determine
stadium duration individuals were maintained in plastic petri dish type con-
tainers, 18 mm high, 80 mm in dia with ventilated tops.
Fifty per cent of the eggs hatched in 7 days, 27% in 8 days and the
remainder in 5 and 6 days (N=97). The duration of the 1st nymphal
stadium was 3-4 days for 70% of the reared individuals while the remainder
290
September, 1983
Baranowski et al.: Euschistus acuminatus
required 5 to 7 days (N=79). The 2nd nymphal stadium required 4 days in
68% of the cases, the remaining individuals ranged from 3-7 days (N=40).
The 3rd instar nymphal stadium also lasted 4 days for most of the individuals
(47%), with the remainder ranging also from 3-7 days (N=38). Half of
the reared individuals required 4 days to complete the 4th nymphal stadium,
and the remainder ranged from 5-9 days (N=34). Twenty individuals were
reared through to adult. The 5th nymphal stadium required 4 days for 1, 6
days for 1, 7 days for 7, 8 days for 6, 9 days for 4, and 1 required 10 days.
Individuals placed on Cestrum nocturnum L., another introduced species
that may also have escaped cultivation, had the same development time as
those on C. diurnum.
One of us (Sailer) has reared several species of Euschistus: servus
(Say), tristigmus (Say), variolarius (P. deB.), ictericus (L.) obscurus
Dall. and latimarginatus Zimmerman on green snap beans (Phaseolus
vulgaris L.). Unlike these species, E. acuminatus apparently will not survive
beyond the 3rd instar on green snap beans. Of 26 newly hatched 1st instars,
22 required 5-6 days to molt, 24-48 hrs longer than individuals on C. diurnum.
Individuals surviving through the 2nd instar also required considerably more
time. Four individuals took 6-9 days and 1 each required 13, 14, and 22 days.
Only 3 individuals survived through the 3rd instar, molting at 5, 8, and 9
days. No individuals survived beyond this stage.
ACKNOWLEDGMENTS
We thank Mary Jane Spring, University of Connecticut, for the prepara-
tion of Figure 1; Mr. E. Pefia and Mr. George Gwinn for collecting ma-
terial; Mr. Jorge Parrado, University of Florida, AREC, for the abstract
translation; and, Ms. Susan Brown for assistance in rearing. This is No.
4414 of the Florida Agricultural Experiment Station Journal Series and
DPI Bureau of Entomology Contribution No. 555.
REFERENCES CITED
BARBER, H. G. 1939. Scientific Survey of Porto Rico and the Virgin Islands:
Insects of Porto Rico and the Virgin Islands: Hemiptera-Heteroptera.
(excepting the Miridae and Corixidae). New York Acad. Sci. 14: 263-
441.
-- AND S. C. BRUNER. 1932. The Cydnidae and Pentatomidae of Cuba.
Jour. Dept. Agr. Puerto Rico 16: 231-84.
BLATCHLEY, W. S. 1926. Heteroptera or true bugs of eastern North America,
with especial reference to the faunas of Indiana and Florida. Indian-
apolis: Nature Publishing Co. 1116 p.
LONG, R. W., AND O. LAKELA. 1971. A flora of tropical Florida. University
of Miami Press, Coral Gables, FL. 962 p.
292 Florida Entomologist 66 (3) September, 1983
THE NORTH AMERICAN PREDACEOUS MIDGES
OF THE BEZZIA BICOLOR GROUP
(DIPTERA: CERATOPOGONIDAE)
WILLIS W. WIRTH
Systematic Entomology Laboratory, IIBIII, Agric. Re?. Serv., USDA,
c/o U.S. National Museum, Washington, DC 20560 USA
ABSTRACT
The bicolor group of species of the predaceous midge genus Bezzia Kieffer,
subgenus Homobezzia Macfie, is diagnosed, and a key is presented for adults
of the 5 North American species, which are described and illustrated. The
North American nominal species B. copiosa (Thomsen) is considered to be a
junior synonym of B. bicolor (Meigen), formerly known only from Eurasia
(NEW SYNONYMY). Bezzia fairchildi NEW SPECIES is described from Florida.
New distribution records are presented for 4 species and the immature stages
are described and figured for 3 species, including detailed larval habitat
records.
RESUME
Se diagnostic el grupo de species bicolor del g6nero de jejenes cerato-
pogonidos predadores Bezzia Keiffer, subgenero Homobezzia Macfie, y se
present una clave para los adults de 5 species norteamericanas, los cuales
se deccriben y se ilustran. La especie norteamericana nominal B. copiosa
(Thomson) se consider ser un sin6nimo junior de B. bicolor (Meigen),
anteriormente conocida solamente de Eurasia (NUEVA SINONOMIA). Bezzia
fairchildi NUEVA ESPECIE se describe de Florida. Se presentan nuevos
registros de la distribuci6n para 4 species, y se described y se ilustran los
estadios inmaduros de 3 species, incluyendo registros detallados de los
habitates larvales.
This review is one of a series in which additions and corrections are made
to the "Revision of the Nearctic Species of the Genus Bezzia" by Dow and
Turner (1976). Former members of Nearctic Bezzia are now recognized as
belonging to 3 genera. Bezzia is now further divided into 2 subgenera and 7
species groups. The primary division of the genus Bezzia into subgenera
Homobezzia Macfie and Bezzia s. str. follows the excellent discussion and
selection of characters by Remm (1974). Remm further divided the group
that I regard as Bezzia s. str. into several additional subgenera, but this
division is not so clear-cut and for the present I prefer to deal with species
groups in the subgenus Bezzia s. str.
In the present series the following have already been studied: Grogan
and Wirth (1981) on the genus Amerohelea Grogan and Wirth; Wirth and
Grogan (1982) on the genus Phaenobezzia Haeselbarth; Wirth (in press)
on the nobilis group in the subgenus Bezzia s. str.; Wirth (in press) on the
cockerelli and dorsasetula groups; and Wirth et al. (in press) on the
annulipes group of the Subgenus Homobezzia. The present paper deals with
another group of the subgenus Homobezzia which is here named the bicolor
group after the earliest described included species, Bezzia bicolor (Meigen).
Wirth: Predaceous Midges 293
A discussion of the taxonomic characters used, and a more complete dis-
cussion of the subgeneric and group classification in the genus Bezzia, are
given in the preceding paper on the Bezzia nobilis Group. The revised classi-
fication as it pertains to North America can conveniently be summarized by
the following key.
SYNOPTIC KEY TO SUBGENERA AND GROUPS OF
NEARCTIC BEZZIA
1. Male antennal segment 12 no longer than 13, antennal plume
weakly developed; (mesonotum dull, occasionally weakly shiny,
brownish or grayish with or without vittae; tibiae pale or with
a dark medial or basal ring; spines of fore femur stout when
present; female with 0.5 pairs of gland rods; males consider-
ably smaller than females; male aedeagus triangular with
minute spinules or hairs) (Subgenus Homobezzia Macfie) ----..- 2
1'. Male antennal segment 12 longest; antennal plume well devel-
oped, extending at least to apex of 13th segment; (mesonotum
black, shiny or dull or with silvery hairs, if grayish brown with
dark vittae, the hind tibia is yellow in midportion, apex broadly
black, and all femora bear spines; tibiae often black; fore
femur with spines slender when present; female abdomen with
1-2 pairs of gland rods; males about same size as female; male
aedeagus variable but not as above) (Subgenus Bezzia Meigen,
s. str.) .....-------....--------- --- ------ 5
Subgenus Homobezzia Macfie
2(1). Larger species, female wing 1.3-3.4 mm long; mesonotum with-
out bristly setae on disc; (fore femur without spines or with
1-4 stout to slender spines of similar lengths, with or without
strong basal tubercles; pupal respiratory horn with numerous
(25-60) spiracular openings, apex more or less flared, abdom-
inal tubercles well developed) --.. ..------- ----........----- 3
2'. Small species, female wing 1.2-1.3 mm long; mesonotum with 2
rows of strong bristly setae on disc; (fore femur with 5-7 stout
ventral spines of alternating uneven lengths arising from dis-
tinct elevations; pupal respiratory horn with only 7-12 spira-
cular openings, abdominal tubercles small) -------- dorsasetula group
3(2). Fore femur unarmed ventrally ----- --- bicolor group
3'. Fore femur armed ventrally with one or more short black spines -.. 4
4(3'). Fore and mid femora entirely dark brown or with dark bands
apical ..-..-..--- ...... ...-------- -- --... ---... --- cockerelli group
4'. Fore and mid femora with subapical dark bands ------ annulipes group
Subgenus Bezzia s. str.
5(1'). Fore femur usually unarmed ventrally; legs brown to black;
femora and tibiae usually with narrow pale rings, rarely
femora pale at base or tibiae pale .-...-..--... --- --_ bivittata group
5'. Fore femur armed ventrally with one or more slender black
spines; legs brown to black, or if banded the pale bands broad -....-. 6
294 Florida Entomologist 66 (3) September, 1983
6(5'). Legs broadly yellow or with broad yellow median bands on fore
femora and tibiae ....--........-.......-.... ----- nobilis group
6'. Legs primarily dark brown to black, at most one pair of legs
with broad yellowish bands -...-----...... ---------....-... expolita group
GENUS Bezzia MEIGEN
SUBGENUS Homobezzia MACFIE
Bezzia bicolor GROUP
DIAGNOSIS. Moderately large pollinose gray midges with brownish mes-
onotal vittae and yellowish scutellum. Wing length 1.3-3.0 mm in female; 0.7-
2.0 mm in male. Vertex without strong median seta. Male flagellum with
distal 5 segments elongated; segment 13 without strong black basal seta. Mes-
onotum (except in coloradensis Wirth) pollinose pale pinkish to dark grayish,
with median brown vitta anteriorly and a pair of lateral vittae posteriorly;
setae of scutellum and above wing bases not strongly developed. Wing trans-
lucent, without markings; anterior veins faintly yellowish. Legs extensively
yellowish, with narrow dark bands usually present, femora and tibiae at most
with broad brown bands on 1 pair of legs; vestiture not strong or bristly;
fore femur unarmed ventrally; female claws small. Female abdomen with 1
or 2 pairs of gland rods. Male basistyles short, dististyles about as long as
basistyles, slightly curved and usually with blunt tip; aedeagus various,
usually triangular; height of basal arch, breadth of distal process, and ex-
tent of ventral spicules vary with species; parameres with long, rod-like
distal process, sometimes slightly narrowed in midportion and/or slightly
broader apically, with tip rounded.
Pupa with respiratory horn long, slender basally and flaring distally,
with 30-60 spiracular openings in a row on distal 1/4 to 1/3; abdominal seg-
ments with strong, thorn-like lateral tubercles, sometimes these are quite
elongate and bearing long setae; posterolateral processes of last segment long
and slender, nearly straight; diverging and densely spinulose. Larva with
head long and slender, nearly cylindrical; body extremely slender, last seg-
ment with long straight anal setae, often as long as segment and directed
more or less anteriorly.
BIOLOGY. The larvae are common inhabitants of mats of filamentous algae
at the water surface of lakes, ponds, and other static water bodies. Chan
and LeRoux (1967) gave a good account of the habits of Bezzia glabra
(Coquillett) in Quebec, where the larvae are predaceous on a variety of
small invertebrates in the algae and may even be cannibalistic. The females,
according to Downes (1978), prey mostly on chironomid midges swarming
near their habitat.
NORTH AMERICAN SPECIES. Bezzia glabra (Coquillett) is widespread over
most of North America and ranges south to Belize; bicolor (Meigen) is
Holarctic and in North America is known from Alaska and Nebraska, and
from Michigan to New York and Connecticut; coloradensis Wirth is es-
sentially Neotropical, known from the Greater Antilles, Florida, and southern
California; fairchildi n. sp. is known only from Florida; and, biannulata
Wirth is known only from California and southern Arizona.
KEY TO THE NEARCTIC SPECIES OF THE Bezzia bicolor GROUP
1. Mesonotum dull waxy black; legs yellowish white, faint brown
Wirth: Predaceous Midges
bands at or near apices of fore and hind femora and at apex of
hind tibia -------------- --- --- coloradensis Wirth
1'. Mesonotum whitish to gray pollinose with brown median vitta;
tibiae with apices blackish and with median pale rings on at
least fore tibia ---- --------. ---------------. 2----------
2. Narrow brown rings at midportion of fore femur and tibia;
apices of all tibiae, hind knee, and base of fore femur brown-
ish .-----.........-...--.....----.. ----------...--- glabra (Coquillett)
Leg markings not as above; all knees dark 3
3. Prominent brown bands at apices of mid and hind femora and
all tibiae and in midportions of all tibiae; proximal third of
hind femur brown .-- ~~-.........-..--...-----------.--- biannulata Wirth
Leg markings not as above; fore femur with median brown
ring and mid tibia without median brown band ..----.......-- ...-...---------.. 4
4. Hind femur brown on proximal 2/3; hind tibia without median
dark band -...- .------- ----...... .------ bicolor (Meigen)
Hind femur pale, at most with faint subapical pale ring; hind
tibia with prominent medial dark band --.... ------------ fairchildi n. sp.
Bezzia biannulata Wirth
(Fig. 1)
Bezzia biannulata Wirth 1952: 237 (male, female; California); Wirth and
Stone 1968: 437 (in key); Cole 1969: 96 (distribution); Dow and
Turner 1976: 123 (redescribed; Figs.; distribution).
DIAGNOSIS. Wing length 2.1-2.5 mm in female; 1.5 mm in male. A mod-
erately large pollinose gray midge with brown mesonotal vittae; legs yellow
with apices of femora, tibiae, and tarsomeres, broad base of hind femur, and
a broad ring in midportion of tibiae black-banded. Female eyes broadly
separated, a distanceof 0.075 mm or 4 facets. Female antenna (Fig. la)
brown, flagellum with narrow bases of segments yellowish; flagellar seg-
ments with lengths in proportion of 25-14-13-13-13-13-13-13-20-20-20-20-20,
antennal ratio 0.85. Male antenna (Fig. Ib) with lengths of segments in
proportion of 25-13-13-13-13-13-12-12-13-16-21-19-22; plume dark brown,
moderately dense, extending to tip of segment 13. Palpus (Fig. Ic) yellow-
ish; lengths of segments in proportion of 5-8-15-10-10; 3rd segment short
and slender with clump of sensilla. Female mandible (Fig. Ig) with 15 mod-
erately large teeth. Thorax dark brown, with dense light gray pollen;
mesonotum (Fig. li) with a pair of narrowly separated brown vittae on
anterior 1/2, and a shorter lateral pair of brown vittae on posterior 1/2; 3
small yellowish setae above wing base; scutellum dull yellowish brown with
4 small setae. Legs (Fig. Ij) yellow; femora slightly brownish toward base,
narrow apex black on fore leg, distal 1/5 black on mid and hind legs, hind
femur with broad basal brown band; tibiae black on distal 1/5, with narrow
brown band at midlength; tarsomeres 1-3 with narrow apices brown, 4-5 all
brown; male legs darker; fore femur without ventral spines; female claws
as in Fig. lh. Wing (Fig. Id) grayish hyaline, anterior veins yellowish;
costal ratio 0.76-0.80 in female, 0.78 in male. Halter infuscated. Female
abdomen dark grayish brown pollinose; 2 pairs of gland rods, the anterior
pair hyaline, the posterior pair brownish; genital sclerotization as in Fig. le,
295
Florida Entomologist 66 (3)
___ A
czx"
e
'Jli \-
m
b
l c--
h
Vk
n\
j/
n
o p
Fig. 1. Bezzia biannulata; a, c-j, female; b, k-1, male; m-q, pupa: a,b,
antenna; c, palpus; d, wing; e, genital sclerotization; f, spermathecae; g,
mandible; h, fifth tarsomere and claws of hind leg; i, mesonotal pattern; j,
color pattern of (top to bottom) fore, mid and hind legs; k, parameres; 1,
genitalia, parameres removed; m, respiratory horn; n, operculum; o (dorsal)
and p (ventral) tubercles of 4th abdominal segment; q, last abdominal seg-
ment.
296
\)----)----------1--7=31,
..-------- .~~_~T~I~3SJ~~
4-y
"Y q
September, 1983
Wirth: Predaceous Midges
a pair of smooth brown areas on 8th sternum. Spermathecae (Fig. If) 2
plus rudimentary 3rd; globular with long necks; subequal, each measuring
0.044 mm in diameter with neck 0.030 mm long. Male abdomen brown;
genitalia (Fig. 11) small; 9th sternum with broad, shallow, caudomedian
excavation; basistyle short and stout; dististyle as long as basistyle, apex
broad and blunt; aedeagus triangular, as long as basal width, basal arch
low, basal arms slender, distal process short and slender with rounded tip.
Parameres (Fig. 1k) with anterolateral arms broad and bilobed; distomedian
process relatively short and stout with rounded tip.
PUPA. About 5 mm long; color light brown with pebble-grained sculptur-
ing. Respiratory horn (Fig. 1m) resembling that of B. glabra, with about
40 spiracular openings on distal 1/3. Operculum (Fig. In) smooth, 3 pairs of
small am tubercles, the anteriormost with a short seta. Abdominal segments
with strong thorn-like tubercles with rudimentary setae as in Fig. lo-p; 2
dasm tubercles located close together, the anterior tubercle nearer midline,
each a small cone with short fine seta; 2 thorn-like 1pm tubercles; dpm3
nearest midline, a low rounded tubercle with rudimentary seta, dpm2
scarcely perceptible just mesad of dpml which is a large, 2-pointed thorn
with a short seta between the 2 points; 2 vpm tubercles arising close to-
gether about 1/2 way between 1pm and midline, each a triangular, flattened
thorn with small subterminal seta. Last abdominal segment (Fig. 1q) with
posterolateral processes nearly straight, long and tapered, about 1/2 again
as long as segment, spinose to tips, not as diverging as those of B. glabra.
LARVA. Unknown.
RECORDED DISTRIBUTION. California.
TYPES. Holotype female, allotype male, Oceano Beach, San Luis Obispo
Co., California, 20-VIII-1948, W. W. Wirth (Type no. 59963, USNM).
NEW RECORD. ARIZONA: Cochise Co., near Portal, Southwest Res. Sta.,
28-V-1967, C. W. Sabrosky, light trap, 1 female.
REMARKS: This species is most closely related to Bezzia bicolor with the
extent of dark leg markings intermediate between those of glabra, coloraden-
sis, and fairchildi on 1 hand and bicolor on the other. The antennal seg-
ments are shorter than those of the first 3 named species, but not as short
as those of bicolor. The configuration of the abdominal tubercles and the
processes of the last abdominal segment of the pupa of biannulata again
are most similar to those of bicolor.
Bezzia bicolor (Meigen)
(Fig. 2)
Ceratopogon bicolor Meigen 1804: 31 (female; Germany); Panzer 1806:
13 (description; fig.); Meigen 1818: 77 (female; Germany; habitat
note); Meigen 1830: 261 (descr. note); Zetterstedt 1850: 3634 (note);
Winnertz 1852: 73 (female descr.; notes); Walker 1856: 234 (notes);
Schiner 1864: 588 (notes; in key; Austria); Gercke 1879: 223 (all
stages; figs.; Germany); de Meijere 1902: 671 (pupal resp. horn; fig.).
Bezzia bicolor (Meigen); Kieffer 1901: 153 (combination; in key); Rieth
1915: 406 (pupa in Spirogyra; fig. resp. horn); Goetghebuer 1920: 105
(male, female redescr.; figs.; Belgium); Goetghebuer 1923: 185 (larva,
pupa; figs.); Edwards 1926: 425 (descr. notes; Britain). Goetghebuer;
in Goetghebuer and Lenz 1934: 77 (redescr.; w. & cent. Europe).
297
Florida Entomologist 66 (3)
... ......
d
September, 1983
eC
Ch
k m
n p rq
Fig. 2. Bezzia bicolor; a, c-k, female; b, 1-m, male; n-r, pupa: a, b, an-
tenna; c, palpus; d, wing; e, mandible; f, genital sclerotization; g,
spermathecae; h, mesonotal pattern; i, anterior view of head; j, fifth
tarsomere and claws of hind leg; k, color pattern of (top to bottom) fore,
mid, and hind legs; 1, parameres; m, genitalia, parameres removed; n,
respiratory horn; o, operculum; p (dorsal) and q (ventral) tubercles of 4th
abdominal segment; r, last abdominal segment.
Probezzia bicolor (Meigen); Kieffer 1919: 129 (combination); Kieffer 1925:
119 (redescr.; figs. larva, pupa); Mayer 1934: 253 (pupa; fig. oper-
culum); Lenz, in Goetbhebuer and Lenz 1984: 123 (pupa in key).
Bezzia (Homobezzia) bicolor (Meigen); Remm 1974: 888 (p. 139 in English
298
I~~----~-~~~
?r ra
Wirth: Predaceous Midges
translation). (in key; descr. notes; USSR distr.; figs.; syn.: typhae
Kieffer 1919 n. syn.; brehmiana Kieffer 1924; algarum Zilahi-Sebess
1930 n. syn.; rufithorax Goetghebuer 1932); Glukhova 1979; 137 (larva;
USSR).
Probezzia typhae Kieffer 1919: 130 (male; Bohemia; figs.); Mayer 1934:
254 (notes on pupa).
Bezzia typhae (Kieffer); Goetghebuer, in Goetghebuer and Lenz 1934: 82
(male redescr.); Lenz, in Goetghebuer and Lenz 1934: 124 (pupa in
key).
Probezzia brehmiana Kieffer 1924: 22 (female: Austria); Mayer 1934:
254 (notes on pupa).
Bezzia brehmiana (Kieffer); Goetghebuer, In: Goetghebuer and Lenz 1934:
78 (female redescr.); Lenz, in Goetghebuer and Lenz 1934: 124 (pupa
in key).
Bezzia brehmii (sic) Kieffer 1925: 130 (in key).
Probezzia algarum Zilahi-Sebess 1930: 197 (all stages; figs.; Hungary);
Zilahi-Sebess 1940: 100 (notes).
Probezzia rufithorax (Goetghebuer 1932: 287 (male, female; Germany);
Mayer 1934: 254 (notes on larva, pupa).
Probezzia copiosa Thomsen 1935: 292 (male, female; New York; figs. );
Thomsen 1937: 79 (larva, pupa; New York; figs.); Johannsen 1943:
785 (in list).
NEW SYNONYMY.
Bezzia copiosa (Thomsen); Johannsen 1952: 167 (combination; in key);
Dow and Turner 1976: 134 (redescr.; figs.; distribution).
DIAGNOSIS. Wing length 1.6-3.0 mm in female, 1.3-2.0 mm in male. A
moderately large, pollinose gray midge species with brown mesonotal vittae;
legs yellow with knees and distal 1/5 of tibiae blackish, femora brown with
subapical pale band, fore tibia with broad median brown band. Eyes (Fig.
2i) broadly separated (a distance of 0.15 mm) in both sexes. Female antenna
(Fig. 2a) short brown with bases of segments pale, very faintly so on seg-
ments 11-15; lengths of flagellar segments in proportion of 15-10-10-10-10-10-
10-10-13-13-13-13-15, antennal ratio 0.76. Male antenna (Fig. 2b) with
lengths of flagellar segments in proportion of 20-12-11-11-10-10-10-10-10-19-
18-19-20; plume brown. Palpus (Fig. 2c) with lengths of segments in propor-
tion of 8-15-21-18-17; 3rd segment short and tapering with clump of sensilla.
Female mandible (Fig. 2e) with 9-13 teeth. Thorax dark brown; mesonotum
(Fig. 2h) with grayish pollen with brown median vitta on anterior 2/3,
with dense brownish pubescence; 3 strong setae above wing base; scutellum
yellowish with 4 marginal setae and dense pubescence. Legs (Fig. 2k)
marked as described above; brownish in male with pale bands often faint;
tarsomeres 1-5 each with narrow apices dark; femora unarmed; female
claws as in Fig. 2j, k. Wing (Fig. 2d) translucent with anterior veins
faintly yellow; costal ratio 0.77 in female,'0.74 in male. Halter pale. Female
abdomen brownish, darker ventrally; 1 pair of yellowish gland rods; genital
sclerotization as in Fig. 2f; 8th sternum with a submedian pair of rounded,
bare, brownish-pigmented areas; 9th sternum a pair of brownish processes
flanking gonopore. Spermathecae (Fig. 2g) 2 plus rudimentary 3rd, heavily
pigmented; ovoid, tapering to moderately long slender necks; unequal,
measuring 0.064 by 0.039 mm and 0.054 by 0.037 mm including necks. Male
299
300 Florida Entomologist 66 (3) September, 1983
genitalia (Fig. 2m) small; 9th sternum with shallow caudomedian excava-
tion; basistyle short and stout, dististyle as long as basistyle, stout to tip;
aedeagus in shape of equilateral triangle basal arch low, scarcely developed,
basal arms slender; ventral surface microspiculose; distal process tapering
to slender tip, sides slightly concave. Parameres (Fig. 21) with anterolateral
processes short, bilobed; basal notch low; distal process moderately stout, of
even breadth throughout, with rounded tip.
PUPA. Brown; length about 4 mm. Respiratory horn (Fig. 2n) slender
proximally, distal 1/2 flaring; 30-40 spiracular openings on distal 1/3.
Operculum (Fig. 20) smooth, with 3 pairs of am tubercles, each with a seta.
Body segments (Fig. 2 p, q) with pebble-grained integument; expanded
laterally at midlength and bearing 3 large, conical 1pm tubercles, the longest
1/2 as long as length of segment, each with a long seta and Ipml also bear-
ing a long subapical seta. Two dasm tubercles, the anterior tubercle located
slightly closer to midline than the other, each with strong rounded base and
long slender seta; 3 dpm tubercles, the 2 laterals contiguous or slightly
joined, elongate and conical with long setae, the mesal tubercle located half-
way between these and midline, very small with minute seta; 2 vpm tubercles,
moderately large, the mesal tubercle with sharp apical point and slender
subapical seta and the lateral tubercle rounded with slender apical seta. Last
abdominal segment (Fig. 2r) very similar to that of B. biannulata, postero-
lateral processes about 1.5 times as long as segment, nearly straight, slender
and tapering to sharp point, spinose to tips, processes not as divergent as
those of B. glabra.
LARVA (from Thomsen, 1937). Length 8-9 mm when mature, color light
green to light tan including head capsule. Head 3 times as long as broad, of
nearly even breadth but slightly tapering at extreme ends; structure and
chaetotaxy as in B. glabra. Last body segment not as slender as in B. glabra,
slightly tapering distally, and there bearing 4 pairs of long, straight, anal
bristles directed anteriorly and nearly as long as segment, plus 2 pairs of
shorter setae 1/4 as long as segment.
NOTE ON SYNONYMY. I have examined a topotypic female and 2 pupal
exuviae of B. copiosa in the USNM collection obtained from Professor
Johannsen, with the following data: "Dryden, N.Y., Ringwood Pond, 9 May
1933, Probezzia copiosa, det. L. C. Thomsen." These agree perfectly with
females in the USNM collection from Monroe Co., New York; Cherry Co.,
Nebraska; and Fairbanks, Alaska, reported by Dow and Turner (1976) as
B. copiosa. They also agree perfectly with the diagnosis and figures of
Remm (1974) for B. bicolor, and with the excellent and detailed description
and figures of B. algarum by Zilahi-Sebess (1930), and with European spec-
imens kindly loaned by the British Museum (Nat. Hist.) from the following
localities:
GREAT BRITAIN: Essex, Epping Forest, 1904, C. Waterhouse, 1 9 and pupal
exuviae. Hants., Sowley Pond, 2-IX-1925, F. W. Edwards, 2 9 2. Herts.,
Radwell, VI-1918, Edwards, 1 9. Inverness, Aviemore, VI-1931, Edwards,
2 9 9. Norfolk, Ormesby, 25-VI-1888, G. H. Verrall, 2 $& 3 9 9. Yorks.,
Austwick, 20-VI-1924, Edwards, 1 9.
RECORDED DISTRIBUTION. "A boreal Transpalaearctic species. Northern
and Central Europe; USSR: Estonia, Lithuania, Ukraine, Karelia, Lenin-
grad, Moscow and Tomsk Provinces, Yakutia, Maritime Territory. Found
Wirth: Predaceous Midges
around various waters (lakes, ponds, bays) and in bogs in July and July"
(Remm 1974). Recorded distribution of B. copiosa in North America: Alaska,
Connecticut, Michigan, Nebraska, New York (Dow and Turner 1976).
NEW RECORDS: ALBERTA: Brooks, 4-7-VIII-1955, J. A. Downes 3 $ 2 9 9
(CNC). MANITOBA: Churchill, 4-VI-1951, 2-29-VII-1953, Downes, 1 13 9 9
(CNC). NORTHWEST TERRITORY: Beaverhill, 62044' N, 104020' W, 29-VI-
1966, G. E. Shewell, 1 9 (CNC); Conrod Lake, 64052/ N, 104015/ W, 8-VII-
1966, Shewell, 2 $ $, 1 9 (CNC). ONTARIO: Point Pelee, 3-VI-1939, Shewell,
1 9 (CNC); Turkey Point, 25-V-1950, J. R. Vockeroth, 1 9 (CNC). WIS-
CONSIN: Dane Co., 28-V-1-VI-1954, R. J. Dicke, light trap, 8 9 9. Washburn
Co., 26-V-1963, R. H. Jones, reared from swamp, 2 S $, 6 9 9, with pupal
exuviae.
DISCUSSION. Adults are quite distinct from the other North American
members of the bicolor group in having the most extensive dark leg mark-
ings and the shortest, distinctly stubby, antennae. The pupa of B. glabra is
easily separable from those of bicolor and biannulata its very elongate ab-
dominal tubercles and diverging posterolateral abdominal processes. The
pupae of B. coloradensis and fairchildi are unknown.
Bezzia coloradensis Wirth
(Fig. 3)
Bezzia coloradensis Wirth, 1952: 238 (male, female; California); Dow and
Turner, 1976: 131 (redescribed; figs.; distribution).
DIAGNOSIS. Wing length 1.3-1.5 mm in female, 1.1 mm in male. A small,
dull-black species with contrasting dull-brown antennae, scutellum, and
abdomen, and yellow halteres and legs. Female eye separation equal to
width of 1 ommatidial facet. Female antenna (Fig. 3a) dull brown, narrow
basses of flagellar segments faintly paler; lengths of flagellar segments in
proportion of 15-10-10-10-10-10-10-10-15-15-15-15-18, antennal ratio 1.10.
Male antenna (Fig. 3b) with lengths of flagellar segments in proportion of
15-10-10-10-10-10-10-10-12-18-22-22-25; segments 4-11 barrel-shaped, 12-15
subcylindrical; plume yellowish, sparse. Palpus (Fig. 3c) short and brown-
ish; lengths of segments in proportion of 6-12-10-8-8; 3rd segment more
slender than 2nd, short, with clump of sensilla. Female mandible (Fig. 3e)
with 10-12 teeth. Thorax dull blackish with rather waxy appearance;
mesonotum with distinct anterior tubercle; 2 fine setae above wing base;
scutellum dull brown, with about 8 microscopic setae. Legs (Fig. 3j) un-
armed, dull dusky yellow with faint brownish bands; bands broad at base of
fore femur, narrow and subapical on all femora, that on mid femur very
faint, narrow at apex of hind tibia; tarsi pale, tarsomere 5 brown; claws as
in Fig. 3i. Wing (Fig. 3d) translucent, anterior veins faintly yellowish;
costal ratio 0.76-0.83 in female, 0.73 in male. Abdomen dull brown; 2 pairs
of yellowish gland rods in female; female genital sclerotization as in Fig. 3f;
8th sternum without perceptible bare areas, 9th sternum with a pair of short,
pointed, brownish processes flanking gonopore. Spermathecae (Fig. 3g) 2
plus rudimentary 3rd, ovoid with tapering necks, darkly pigmented; unequal,
measuring 0.061 by 0.037 mm and 0.046 by 0.027 mm including necks. Male
genitalia (Fig. 3k) small; 9th sternum with very shallow caudomedian
excavation; basistyle stout at base; dististyle about as long as basistvle.
301
Florida Entomologist 66(3)
September, 1983
a
,,.. .-. .. ..... .
d
e
f h
Fig. 3. Bezzia coloradensis; a, c-h, j, female; b, i, k, male: a, b, antenna;
c, palpus; d, wing; e, mandible; f, genital sclerotization; g, spermathecae;
h, fifth tarsomere and claws of hind leg; i, parameres; j, color pattern of
(top to bottom) fore, mid and hind legs; k, genitalia, parameres removed.
tapering to slender tip; aedeagus elongate, twice as long as basal breadth,
only slightly concave on proximal margin, sides nearly straight, tapering to
slender tip which is slightly truncated. Parameres (Fig. 3i) with short,
bilobed anterolateral arms, distinct anteromedian notch; distomedian process
elongate, tapering to slender midportion and gradually expanded distally to
a spatulate, rounded tip.
IMMATURE STAGES. Unknown.
RECORDED DISTRIBUTION. California, Florida.
TYPES. Holotype female, allotype male, Westmorland, Imperial Co., Cali-
fornia, 6-IV-1949, W. W. Wirth, swept from alkaline slough (Type no. 59963,
USNM).
NEW RECORDS. CAYMAN ISLAND: Grand Cayman, XII-1973, J. E. Davies,
10 9 9. HAITI: Chou Chou Baie, 8-VI-1978, C. Raccurt and R. Lowrie, swept,
4 9 9. JAMAICA: St. Catherine Parish, Twickenham Park, 28-IV-1970, E. G.
Farnworth, light trap, 1 9. VIRGIN ISLANDS: St. John, 9-12-IX-1961, R. W.
Williams, emergence trap, 46 9 9.
DISCUSSION. This species differs markedly from the other North American
302
Wirth: Predaceous Midges
303
species of the bicolor group in its dull blackish mesonotum and quite re-
stricted dark bands on the legs. Its essentially Neotropical distribution cor-
responds with its somewhat peripheral taxonomic position in the group.
Bezzia fairchildi Wirth, NEW SPECIES
(Fig. 4)
DIAGNOSIS. A medium-sized brown and yellow midge with pollinose gray
thorax with brown median vitta on mesonotum; legs with middle of fore
and hind tibiae, and ends of leg segments narrowly brown-ringed.
HOLOTYPE FEMALE. Wing length 1.74 mm; breadth 0.67 mm.
Head: Brown, narrow bands of antennal flagellar segments yellowish.
Eyes separated by width of 1 ommatidial facet. Antenna (Fig. 4a) with
lengths of flagellar segments in proportion of 15-11-10-10-10-10-10-11-30-29-
28-29-30, antennal ratio 1.68. Palpus (Fig. 4b) with lengths of segments in
proportion of 5-12-16-13-12; 3rd segment slender, with clump of sensilla,
palpal ratio 3.2. Mandible with 10 teeth.
Thorax: Brown; mesonotum pollinose gray with median brown vitta on
anterior 1/2 and lateral brown vittae above wing bases; scutellum yellowish
with 4 yellowish marginal setae. Legs (Fig. 4h) yellow; coxae and tro-
chanters brown; fore and hind knees blackish; apices of tibiae and median
bands on fore and hind tibiae blackish; sometimes faint brownish bands
present subapically on femora and in middle of mid tibia; narrow apices of
tarsomeres brown; femora unarmed; claws (Fig. 4g) large, gently curved;
each 0.07 mm long on fore and mid legs, 0.10 mm long on hind leg, each with
minute inner basal tooth. Wing (Fig. 4c) translucent with anterior veins
creamy white; costal ratio 0.83. Halter pale.
Abdomen: Pale brown dorsally, yellowish ventrally; 2 pairs of brownish
gland rods. Genital sclerotization as in Fig. 4e. Spermathecae (Fig. 4f) 2
plus rudimentary 3rd; oval with short neck scarcely evident; heavily pig-
mented, dark brown; unequal, measuring 0.078 by 0.058 mm and 0.075 by
0.043 mm.
ALLOTYPE MALE. Wing length 1.53 mm; breadth 0.61 mm. Similar to the
female, with the usual sexual differences. Thorax and abdomen dull brown-
ish, the brownish bands on the legs very indistinct. Antenna with lengths of
flagellar segments in proportion of 23-14-13-13-13-13-13-15-23-33-30-30-32,
antennal ratio 1.25; plume sparse, pale yellowish. Costal ratio 0.77. Genitalia
(Fig. 4 i-j) nearly identical with those of B. glabra, with the following most
important differences: shorter and broader, the convex lateral margins of
the basistyles and dististyles forming a nearly circular outline; ninth
sternum with caudomedian excavation forming a semicircle, extending half-
way to base of segment.
DISTRIBUTION. Florida.
TYPES. Holotype female, Blackwater River Biological Station, Santa
Rosa Co., Florida, 21-V-1971, G. B. Fairchild, light trap (Type no. 76594,
USNM). Allotype male, Leon Co., Florida, Tall Timbers Research Station,
IV-1970, F. W. Mead, light trap. Paratypes, 10 8$ 85 9 9, as follows:
FLORIDA: Alachua Co., Gainesville, Chantilly Acres, IV-V-1967, F. S.
Blanton, light trap, 3 9 ; same, 18-IV-1967, W. W. Wirth, Malaise trap,
1 9; West Gainesville, V-1968, Blanton and Fletcher, light trap, 2 9 9;
Florida Entomologist 66 (3)
September, 1983
b
C
_* ,,,^*"
e .
304
a
Vd
k
0
Fig. 4. Bezzia fairchildi; a-g, female; h-i, male: a, antenna; b, palpus;
c, wing; d, mandible; e, color pattern of (top to bottom) fore, mid and hind
legs; f, genital sclerotization; g, spermathecae; h, genitalia, parameres re-
moved; i, parameres.
Hawthorne, 27-IV-1968, Blanton, light trap, 1 Q ; 3.5 mi W Lochloosa, 19-IV-
1967, Blanton, light trap, 13 9 9. Glades Co., Palmdale, 14-VII-1970, E. Irons,
light trap, 1 9. Gulf Co., 2 mi N Beacon Hill, 15-X-1970, Blanton, light trap,
1 9. Highlands Co., Lake Placid, 1-7-V-1964, R. W. Hodges, light trap, 1 9 ;
same, 19-IV-1970, J. N. Layne, light trap, 3 9 9 ; Sebring, 8-VIII, 15-X-1942,
C. T. Parsons, 2 9 Y (MCZ). Hillsoboro Co., Harris Swamp, 13-IV-1967,
Blanton, light trap, 11 9 9. Leon Co., Tall Timbers Res. Station, IV-1970,
F. W. Meade, light trap, 3 S $, 11 9 9 ; 29-V-1973, Wirth, light trap, 1 9 ;
3 mi N Tallahassee, V-1970, Blanton, light trap, 3 S S, 11 9 9. Liberty Co.,
Torreya St. Park, 22-IV-1967, Wirth, Malaise trap, 1 9. Marion Co., Juniper
Springs, 28-IV-1970, Wirth, light trap, 1 '9 Orange Co., Rock Springs, 21-IV-
1970, Wirth, light trap, 1 9. Putnam Co., Lon's Lake, V-VI-1971, Blanton,
light trap, 4 8 $, 7 9 9; Welaka, 9-IV-1964, H. A. Denmark, light trap, 1 9.
Santa Rosa Co., Blackwater River Biological Sta., 21-V-1971, Fairchild, light
trap, 10 9 9 ; 25-V-1973, Wirth, light trap, 2 9 9. Wakulla Co., Ochlockonee
River St. Park, 29-IV-1970, Wirth, light trap, 1 9. Paratypes deposited in
BMNH, CAS, CNC, USNM.
DISCUSSION. This species is named for G. B. Fairchild of the University
f
I
Wirth: Predaceous Midges
of Florida in appreciation of his friendship and in recognition of his im-
portant contributions to our knowledge of Florida and Neotropical Diptera.
Bezzia fairchildi is closely related to B. glabra but can readily be dis-
tinguished by the prominent median brown band on the hind tibia.
Bezzia glabra (Coquillett)
(Fig. 5)
Ceratopogon glaber Coquillett 1902: 85 (female; Florida).
Bezzia glabra (Coquillett); Kieffer 1906: 58 (combination); Wirth 1952:
236 (redescribed; figs.; all stages; California); Johannsen 1952: 167
(in key; fig. aedeagus); Chan and LeRoux 1967: 16 (larval biology;
Quebec); Dow and Turner 1976: 145 (redescribed; figs.; distribution);
Downes 1978: 1-62 (female prey records; Canada); Palchick 1981: 266
(pupa; figs.; Wisconsin).
Probezzia glabra (Coquillett); Johannsen 1908: 267 (combination); Mal-
loch 1914: 137 (in key); Malloch 1915: 355 (notes; Illinois; fig. wing);
Muttkowski 1918: 407 (notes on larva; Wisconsin); Thomsen 1935:
293 (male; New York; fig. aedeagus); Thomsen 1937: 78 (immature
stages; New York; biology; figs.); Peterson 1951: 266 (larva; figs.).
DIAGNOSIS. Wing length 1.3-2.4 mm in female, 0.7-1.6 mm in male. A
small to medium-sized pale grayish midge with broad reddish-brown
mesonotal vitta; darker in male. Female eyes (Fig. 5d) separated by width
of 2 ommatidial facets. Female antenna (Fig. 5a) pale brown with narrow
bases of flagellar segments whitish; lengths of flagellar segments in propor-
tion of 15-9-8-8-8-8-8-9-26-27-24-27-25, antennal ratio 1.3-1.8. Male antenna
with sparse yellowish plume. Palpus (Fig. 5b) with lengths of segments in
proportion of 10-14-30-20-20; palpal ratio 2.1-4.0. Female mandible with
11-16 coarse teeth. Thorax pearly pinkish gray pollinose; mesonotum with
broad median reddish-brown vitta on anterior 2/3 plus a pair of brownish
lines on sides; scutellum shining yellowish brown with about 8 small yellow-
ish setae; usually 3 coarse setae above wing base. Legs (Fig. 5e) yellowish,
narrow dark brown rings on fore and hind knees, ends of all tibiae, and
fainter at midlengths of fore femur and tibia; all tarsomeres narrowly dark
at apices; hind basitarsus elongate, TR 2.3-3.3; claws large, gently curved;
0.06 mm long on fore and mid legs, 0.08 mm long on hind leg, each claw with
minute inner basal tooth. Wing (Fig. 5c) translucent with anterior veins
yellowish white; costa long, female costal ratio 0.75-0.80, in male 0.63-0.78.
Halter pale. Female abdomen dull whitish to pale brownish; 1 pair of yellow-
ish gland rods as long as 4.5 segments; genital sclerotization (Fig. 5f) with
2 brownish pigmented areas on setose lobes of 8th sternum; 9th sternum a
pair of prominent dark brown processes flanking gonopore. Spermathecae
(Fig. 5f) 2 plus rudimentary 3rd; elongate oval with short slender neck;
darkly pigmented, blackish; slightly unequal, measuring 0.108 by 0.059 mm
and 0.106 by 0.054 mm. Male abdomen brownish; genitalia (Fig. 5g) small,
short; 9th sternum with deep caudomedian excavation; basistyle slightly
tapering, a low rounded setose basimedian lobe; dististyle nearly as long as
basistyle, tapering to bluntly pointed tip; aedeagus a very short, triangular,
sclerotized plate with concave anterior arch about 1/3 total length.
Parameres with strongly sclerotized, stout, basilateral lobes; distal process
305
306 Florida Entomologist 66 (3) September, 1983
a
iii
SC O
Sr d
ki
h \ ... / ."
i.j ii / m n.
k 0
Fig. 5. Bezzia glabra; a-f, female; g', male; h-j, larva; k-o, pupa: a,
antenna; b, palpus; c, wing; d, anterior view of head; e, hind, med, and fore
legs (left to right); f, genital sclerotization and spermathecae; g, genitalia;
h, head and anterior thoracic segments, dorsal view; i, head, ventral view;
j, last abdominal segment; k, respiratory horn; 1, operculum; m (ventral)
and n (dorsal) tubercles of 4th abdominal segment; o, last abdominal seg-
ment. (a-g, m-o drawn by Ethel Grogan; 5h-1 from Wirth, 1952).
Wirth: Predaceous Midges
long and slender with slightly expanded, rounded tip, slightly expanded near
base and narrowed in midportion.
PUPA. Yellowish, about 4.0 mm long. Respiratory horn (Fig. 5k) long
(0.043 mm in male, 0.050 mm in female), slender at base and gradually
flaring to darkened tip 3 times basal breadth, bearing 50 (male) to 60
(female) spiracular openings. Operculum (Fig. 51) without sexual dimor-
phism, 1.5 times as wide as long, 3 pairs of am tubercles, the 2 anteriormost
each with a fine seta. Abdominal segments (Fig. 5 m-n) with 3 long 1pm
tubercles ending in a long seta, Ipml tubercle 0.070 mm long, with an
additional long seta arising at about 0.3 way from base; 2 dasm tubercles,
nearly in longitudinal line, each with long seta; 3 conical dpm tubercles, each
with a long seta, tubercles dpml and dpm2 fused; the 2 pairs of dasm
tubercles and the pair of dpm3 tubercles forming on each segment an al-
most equal-sided hexagonal pattern of similar tubercles; 3 conical vpm
tubercles, each with a long seta, vpml and vpm2 fused in a common tubercle.
Last segment (Fig. 5o) with posterolateral processes long and slender, about
twice as long as segment, straight and slightly divergent, spinose nearly to
trips.
LARVA. Very slender, length about 10 mm when mature; color creamy
whitish, head yellowish. Head (Fig. 5 h,i) about 4 times as long as broad,
of nearly even breadth, slightly tapering at both ends. Last body segment
(Fig. 5j) about 10 times as long as broad, apex with ring formed by 4 pairs
of long dark-brown straight hairs, directed somewhat anteriorly, each about
0.7 as long as segment; also 2 pairs of shorLer setae.
DISTRIBUTION: North American from Alaska to California and southern
Ontario, Quebec, and New Brunswick to Florida; also Belize.
TYPES: Two female syntypes, Biscayne Bay, Florida (Mrs. Annie T.
Slosson) (Type no. 6155, USNM). Lectotype female, pinned, the specimen
bearing Coquillett's determination label, by present designation.
SPECIMENS EXAMINED. Pinned, 182 specimens; slides, 83 S 8, 138 9 9.
NEW NEOTROPICAL RECORD. BELIZE: Cayo Dist., Western Hwy MP 66,
VI-1969, W. and D. Haase, light trap, 6 9 2.
RECORDS OF REARED MATERIAL. CALIFORNIA: Colusa Co., Grimes, 22-IX-
1965, rice field (Washino), 1 S with pupal exuviae. Fresno Co., Selma,
Rockwell Pond, 4-VIII-1947 (Wirth), all stages, many specimens. Los
Angeles Co., Malibu Lake, 5-IX-1951 (Belkin), reared, 2 pupae. LOUISIANA:
Baton Rouge, University Lake, 25-II-1947 (Wirth), algal mat, 10 pupae.
MARYLAND: Montgomery Co., Fairland, 27-V-1959 (Hubert), reared, 1 S
and pupal exuviae. Pince George's Co., College Park, Lakeland Pond, 26-
V-1975 (Grogan), reared, 4 & S with pupal exuviae; Patuxent Wildlife
Res. Center, 29-VI-1976 (Grogan), ex pond weeds, 6 larvae. QUEBEC: St.
Anne de Bellevue, Morgan Arboretum, ex pond algae, 1964 (Chan), 4 larvae,
3 pupae. VIRGINIA: Augusta Co., Mount Solon, 12-VI-1950 (Wirth), 2 pupae.
ACKNOWLEDGEMENTS
I am very grateful to J. Antony Downes of the Biosystematics Research
Institute, Agriculture Canada, Ottawa; William L. Grogan, Jr., of Salisbury
State College, Salisbury, Maryland; Walter I. Knausenberger of the College
of the Virgin Islands, St. Croix; and Susan M. Palchick of the University of
California at Davis, for critical reviews of the manuscript and many helpful
307
Florida Entomologist 66 (3)
comments and suggestions. I wish to acknowledge with thanks the loan of
specimens from the following collections: British Museum (Nat. Hist.),
London (BMNH), courtesy of R. Lane; California Academy of Science, San
Francisco (CAS) through P. H. Arnaud, Jr.; Canadian National Collection,
Ottawa (CNC) through J. A. Downes; Cornell University, Ithaca, New York
through Q. D. Wheeler; Illinois Natural History Survey, Urbana through
D. W. Webb; Museum of Comparative Zoology, Cambridge, Massachusetts
through Karen Jepson; University of California, Insect Survey, Berkeley
through D. Wagner; and University of California, Riverside through Saul
Frommer. Ms. Molly K. Ryan prepared the illustrations.
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midges (Diptera: Ceratopogonidae) in Quebec. Ann. Soc. Ent. Quebec
12: 14-68.
COLE, F. R. 1969. The Flies of Western North America. Univ. California
Press, Berkeley. 693 p.
COQUILLETT, D. W. 1902. New Diptera from North America. Proc. United
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Dow, M. I., AND E. C. TURNER, JR. 1976. A taxonomic revision of the
Nearctic species of the genus Bezzia (Diptera: Ceratopogonidae). Res.
Div. Bull. Virginia Polytech. Inst. St. Univ. 103: 1-162.
DOWNES, J. A. 1978. Feeding and mating in the insectivorous Ceratopo-
goninae (Diptera). Mem. Ent. Soc. Canada 104: 1-62.
EDWARDS, F. W. 1926. On the British biting midges (Diptera, Ceratopo-
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GERCKE, G. 1879. Ueber die Metamorphose nacktfluigeliger Ceratopogon-
Arten sowie fiber die von Tanypus nigropunctatus Steg. und von
Hydrellia mutata Meig. Verhandl. Ver. naturw. Unterhalt. Hamburg
4: 222-8, 1 plate.
GLUKHOVA, V. M. 1979. Larvae of biting midges of the subfamilies Pal-
pomyiinae and Ceratopogoninae of the fauna of the USSR (Diptera,
Ceratopogonidae = Heleidae) (In Russian). Akad. Nauk SSSR Izdv.
Zool. Inst. Faune SSSR no. 121: 1-230. Leningrad.
GOETGHEBUER, M. 1920. Ceratopogoninae de Belgique. Mem. Mus. Roy. Hist.
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1923. Larves et nymphes de C6ratopogonines. Ann. Biol. Lacustre
11: 173-86.
.1932. Ceratopogonidae et Chironomidae nouveaux ou peu connus
d'Europe. Deuxieme Note (1). Bull. Ann. Soc. Ent. Belgium 72:
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GOETGHEBUER, M., AND F. LENZ. 1933-4. Heleidae (Ceratopogonidae). Fam.
13a, Pages 1-48, plates 1-6 (1933); Pages 49-133, plates 7-12 (1934).
In: E. Lindner, Ed. Die Fliegen der Palaearktischen Region. Vol. 3.
Stuttgart.
GROGAN, W. D., JR., AND W. W. WIRTH. 1981. A new American genus of
predaceous midges related to Palpomyia and Bezzia (Diptera:
Ceratopogonidae). Proc. Biol. Soc. Washington 94: 1279-305.
JOHANNSEN, O. A. 1908. New North American Chironomidae. New York
St. Mus. Bull. 124: 264-85.
1943. A generic synopsis of the Ceratopogonidae (Heleidae) of the
Americas, a bibliography, and a list of the North American species.
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_. 1952. Guide to the insects of Connecticut. Part VI. The Diptera or
September, 1983
308
Wirth: Predaceous Midges 309
true flies of Connecticut. Fifth fascicle: Midges and gnats (part).
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KIEFFER, J. J. 1901. Synopse des representants europ6ens du group
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1919. Chironomides d'Europe conserves au Mus6e National Hongrois
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1924. Chironomides nouveaux ou rares de l'Europe central. Bull.
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1925. DiptBres (Nematocres piqueurs): Chironomidae Ceratopo-
goninae. Faune de France. 11: 1-139. Paris.
MALLOCH, J. R. 1914. Synopsis of North American species of the genus
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1915. The Chironomidae or midges of Illinois. Bull. Illinois St. Lab.
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MAYER, K. 1934. Die Metamorphose der Ceratopogonidae (Dipt.). Arch. f.
Naturges. 3: 205-88.
MEIGEN, J. W. 1804. Klassifikazion und Beschreibung der europaischen
zweifliigeligen Insecten (Diptera Linn.). 1: 1-314, 15 plates. Braun-
schweig.
1818. Bystematische Beschreigung der bekannten europaischen
zweifliigeligen Insekten. 1: 1-333, 11 plates.
1830. Systematische Beschreibung der bekannten europaischen
zweifliigeligen Insekten. 6: 1-401, 11 plates.
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Spuppen. Zool. Jahrb. Abth. f. Morph. 15: 623-92, 4 plates.
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and pupae of six species of Bezzia and three species of Nilobezzia
(Diptera: Ceratopogonidae). M.S. Thesis, Univ. Wisconsin, Madison.
28 p., 12 plates.
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310 Florida Entomologist 66 (3) September, 1983
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predaceous midges of the Bezzia annulipes Group (Diptera: Ceratopo-
gonidae). Proc. Ent. Soc. Washington (In Press)
WIRTH, W. W., AND A. STONE. 1968. Aquatic Diptera. Pages 372-482. In:
Usinger, R. L., Ed. Aquatic Insects of California. Univ. California
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ZETTERSTEDT, J. W. 1850. Diptera Scandinaviae. Disposita et descripta. 9:
3367-710.
ZILAHI-SEBESS, G. 1930. Zwei neue Chironomidenarten aus dem Balaton-
gebiet. Kozl. Tesza Inst. Debrecen 6: 197-205, 2 plates.
1940. Magyarorszag Heleidai. Folia Ent. Hungarica 5: 1-136, 3
plates.
REDUCED EFFICACY OF FENBUTATIN-OXIDE IN
COMBINATION WITH PETROLEUM OIL IN
CONTROLLING THE CITRUS RUST MITE
PHYLLOCOPTRUTA OLEIVORA1
C. C. CHILDERS AND A. G. SELHIME2
University of Florida, IFAS
Agricultural Research and Education Center
Lake Alfred, Florida 33850
ABSTRACT
Fenbutatin-oxide, Vendex, was applied as a 50% wettable powder or
4L liquid formulation alone and in combination with either medium (FC-435)
or light (FC-412) petroleum oils on citrus to control the citrus rust mite,
Phyllocoptruta oleivora (Ashmead). Lower rates of the miticide in com-
bination with either oil at 1% resulted in a significant reduction in residual
activity in controlling rust mite versus the miticide applied alone.
RESUME
Se aplicaron a citrus para el control de Phyllocoptruta oleivora (Ash-
mead) el acaricida VendexR en forma de polvo mojable de 50%, o en la
formulacion liquid 4L solo y en combinaci6n con petroleo de peso o medio
(FC-435) o ligero (FC-412). En comparasi6n con el acaricida solo, las con-
centraciones bajas del acaricida en combinaci6n con qualquier de los petroleos
'Acari: Eriophyidae.
2Horticultural Research Laboratory, USDA, ARS, Orlando, FL 32803.
Childers & Selhime: Citrus Rust Mite Control
en una concentraci6n de 1% result en una rebajada significativa de la
actividad residual en control del Acaro.
Field trials to evaluate the efficacy of fenbutatin-oxide, Vendex, alone
and in combination with petroleum oil to control the citrus rust mite (CRM),
Phyllocoptruta oleivora (Ashmead) were conducted between 1980 and 1982
in central Florida. The objective of these trials was to determine efficacy of
the miticide at lower than recommended rates and to determine if petroleum
oil tank mixed with Vendex would provide extended control of CRM. The
recommended rate of Vendex in the 1983 Florida Citrus Spray Guide is 2 to
4 oz ai/100 gal (Knapp 1983).
Medium and light petroleum oils (FC-435 and FC-412, respectively) are
commonly used on Florida citrus in spray-tank mixes. Usually, the summer
spray applied in June to July includes oil to control the fungal disease, greasy
spot, Mycosphaerella citri Whiteside, to enhance insect and mite control, and
to loosen sooty mold accumulation on leaves (Simanton and Trammel 1966).
This paper reports on the effectiveness of 2 formulations of Vendex for
CRM control when applied alone or in combination with either oil formula-
tion. Additional experiments evaluating Vendex alone and/or in combina-
tion with oil are presented in Insecticide and Acaricide Tests (Childers and
Konsler 1981, Childers and Sorrell 1982, 1983).
MATERIALS AND METHODS
On 3 June 1980, Vendex 50 WP was applied at rates of 1, 2 and 3 oz
ai/100 gal alone and in combination with 1% FC-435 oil in a young 'Hamlin'
orange grove in the vicinity of Montverde, FL. Other treatments were ethion
4EC at 6 oz ai/100 gal and a water sprayed check. Spray plots consisted of
single trees replicated 7 times. The materials were applied dilute to runoff
at 350 psi using a hydraulic sprayer equipped with a handgun. Tree height
was ca. 7 ft with a 6 ft diam and each tree received ca. 5 gal of spray. The
water pH was 6.1.
Treatments were assigned to single-tree plots in a complete randomized
block design based on total pretreatment counts/tree of living motile citrus
rust mites. The highest counts were assigned to the first replicate. The next
series of counts were assigned to the second replicate followed eventually by
the lowest mite counts assigned to the last replicate. Randomization of treat-
ment assignment to trees within each replicate followed.
Twenty fruit were examined at random around the canopy of each treated
tree at various times before and after spraying using a 10X handlens with
an attached 1 cm2 grid area patterned after Allen (1976) (Fig. 1). A 1 cm2
area was examined on each of 2 shaded sides per fruit. Population density
estimates of CRM were obtained by counting the number of living motile
mites present within the combined 2 cm2 count per fruit and recorded as 1
observation. Means in tables are based on 2 observations per fruit with a
total of 20 fruit per replicate. The number of CRM per observation was
transformed by VX + 0.5 for statistical analysis. Non-transformed means
are presented in the accompanying tables.
In 1981, Vendex 4L treatments were applied to a young, mature 'Wash-
ington' navel orange grove at Dundee, FL. The only additional treatment
Florida Entomologist 66 (3)
Fig. 1. 10X handlens equipped with 1 cm2 grid used in determining citrus
rust mite densities on fruit surfaces.
was an untreated check. Spray plots consisted of 4 trees replicated 5 times
arranged in a randomized block design. Five fruit were examined at random
around the canopy of each of 4 sample trees or 20 fruit/replicate at various
times before and after spraying. Mite numbers were evaluated as in the
1980 test. The materials were applied dilute to runoff at 400 psi using a
hydraulic sprayer equipped with a handgun. Trees were 7 ft with ca. 7 ft
diam and each received ca. 7 gal of spray. Water pH was 7.2.
A second test with Vendex 4L was applied in 1981 with the same treat-
ment sequence and numbers of fruit sampled for mites as in the 1980 test.
Treatments were applied to single tree plots replicated 5 times as described
in 1980 at 550 psi in a mature 'Ruby' grapefruit block near Avon Park, FL.
Tree height was 8-10 ft with ca. 13 ft canopy diam. Each tree received ca.
8 gal of spray and water pH was 8.1.
In 1982, Vendex 4L at 1.75 oz ai/100 gal was applied on 3 May alone and
in combination with light oil, basic copper sulfate, or Dithane Z-78 in a
mature 'Hamlin' orange grove near Groveland, FL. This approximates the
recommended label rate of 2-1/2 pints of Vendex per acre. Treatment assign-
ment and fruit sampled for mites were the same as in the 1980 test. About
312
September, 1983
Childers & Selhime: Citrus Rust Mite Control
8 gal of spray per tree were applied dilute at 400 psi and treatments were
replicated 6 times. Tree height was 13 ft with 15 ft canopy diam.
Percentage of fruit area with visible CRM injury and percentage of
rejected fruit below U.S. Fancy standards were determined. Fifty fruit were
selected at random around the canopy perimeter of each sample tree on the
indicated date(s) and the percentage of surface russetted due to CRM feed-
ing was estimated per fruit. Data were subjected to arcsine VY transforma-
tions for statistical analysis.
Mite-days were determined for these same 2 experiments by calculating
the area under the population growth curve over time (Allen 1976). Mite-
days were calculated using the formula
D + D2 SI
2
where D1 and D2 represent the mite densities at times 1 and 2 and SI is the
sample interval expressed in days. Intervals are additive to provide cumu-
lative mite-days.
RESULTS
Reduced residual control of CRM by Vendex 50 WP at 1 and 2 oz ai/100
gal in combination with 1% FC-435 oil occurred by 34 days postspray in the
1980 test with control lost by 52 days (Table 1). The Vendex + oil combina-
tions provided comparable control of CRM to ethion at 6 oz ai, however,
Vendex alone resulted in superior residual control.
In 1981, similar results occurred in the first test (Table 2). Vendex 4L
at 1 oz ai/100 gal + 1% FC-435 oil began losing residual activity by 42 days
postspray. A significant difference in mite control occurred by 56 days post-
spray at the 1 and 2 oz ai/100 gal of Vendex rates in combination with oil
versus Vendex alone. A significant difference in mite control was evident
between the 3 oz ai rate of Vendex alone versus the corresponding rate com-
bined with oil by 70 days.
A second test was conducted in 1981 at Avon Park. Again, the low rate
of Vendex when combined with 1% medium oil was significantly less effective
by 15 days postspray. Control was lost by 39 days (Table 3). Higher but
non-significant rust mite densities were obtained at the 2 and 3 oz ai rates of
Vendex combined with oil versus Vendex alone. Higher but non-significant
differences in percentage of rejected fruit were obtained for the miticide-oil
combinations.
The experiment was terminated at this critical time when the grower
insisted upon spraying for CRM control when unacceptable levels of rus-
seting developed on fruit in the buffer trees in the block. The 'Ruby' grape-
fruit in this block were being grown for fresh market. The entire block was
sprayed with sulfur on 1 September and excellent mite control was achieved.
In 1982, no significant differences were obtained during the brief period
of rust mite buildup (ca. 30 days) as indicated by mite density counts and
total cumulative mite-days (Table 4). However, a significantly greater
percentage of rejected fruit in the Vendex + 1% light oil combination oc-
curred. A paired comparison of Vendex alone versus Vendex + 1% light oil
treatments using analysis of variance and Fisher's LSD resulted in F-values
313
Florida Entomologist 66 (3)
September, 1983
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Childers & Selhime: Citrus Rust Mite Control 317
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Florida Entomologist 66 (3)
of 5.95 (1, 10 df) for % russeted fruit and 5.96 for % rejected fruit. Both
were significant at the 5% level. Comparable paired comparisons of Vendex
versus Vendex + 0.5% FC-412 oil were not significantly different.
DISCUSSION
Vendex versus Vendex + oil comparisons can only be evaluated effectively
when rust mite population pressure is sustained as indicated by increasing
population density readings over time, particularly 30 to 60 days postspray.
This is not always obtainable since population collapse often occurs from
the fungal pathogen, Hirsutella thompsonii Fisher (Fisher 1950) or un-
favorably hot-dry microclimatic conditions (Reed et al. 1964, Hobza and
Jeppson 1974). Despite these impediments, results of several field trials have
demonstrated differences between residual efficacy of lower rates of Vendex
alone versus Vendex in combination with 1% FC-435 or FC-412 oil. Virtually
all growers would use 1% or more oil in tank mixes for concentrate applica-
tion on citrus. Other miticides including amitraz, carbosulfan (Childers,
unpub data) and chlorobenzilate (R. Johnson, unpub data) have been evalu-
ated against CRM in combination with 1% petroleum oil without a reduction
in residual control versus the miticides applied alone.
Whether reduction in activity of Vendex with oil is the result of a
physical interference or a chemical reaction has not been determined. Possi-
ble antagonism could occur by the oil engulfing the miticide and thus pre-
venting contact with the target mite species. Conversely, a chemical reaction
may occur which hastens deactivation of miticidal properties of the com-
pound.
Vendex is an excellent miticide against the CRM; the Texas citrus mite,
Eutetranychus banksi (McGregor); and the citrus red mite, Panonychus
citri (McGregor) in Florida. The miticide has been found to be selective in
its control of pest mites while minimizing mortality of predaceous arthropods
(Jeppson et al. 1975, Asquith and Hull 1979). However, due to its relative
high cost and loss of residual activity when mixed with 1% oil, this miticide
would not be a material of choice for use in tank mixes containing oil on
citrus.
ENDNOTE
This paper reports the results of research only. Mention of a pesticide in
this paper does not constitute a guarantee or warranty by the U.S. Depart-
ment of Agriculture or the University of Florida nor does it imply registra-
tion under FIFRA.
ACKNOWLEDGEMENTS
Florida Agricultural Experiment Stations Journal Series No. 4593. We
wish to thank Mr. R. W. Sorrell and Mr. B. A. Konsler of the AREC, Lake
Alfred for technical assistance.
REFERENCES CITED
ALLEN, J. C. 1976. A model for predicting citrus rust mite damage on
Valencia orange fruit. Environ. Ent. 5 (6) : 1083-8.
318
September, 1983
Childers & Selhime: Citrus Rust Mite Control
ASQUITH, D., AND L. A. HULL. 1979. Integrated pest management systems
in Pennsylvania apple orchards. Pages 203-22. In: D. J. Boethel and
R. D. Eikenbary, eds. Pest Management Programs for Deciduous Tree
Fruits and Nuts. Plenum Press. New York. 256 p.
CHILDERS, C. C., AND B. A. KONSLER. 1981. Chemical control of citrus rust
mite on citrus. Insecticide and Acaricide Tests 6: 44.
AND R. W. SORRELL. 1982. Chemical control of citrus rust mite on
citrus. Insecticide and Acaricide Tests 7: 56-8.
AND 1983. Chemical control of citrus rust mite on citrus.
Insecticide and Acaricide Tests (In press).
FISHER, F. E. 1950. Entomogenous fungi attacking scale insects and rust
mites on citrus in Florida. J. Econ. Ent. 43 (3) : 305-9.
HOBZA, R. F., AND L. R. JEPPSON. 1974. A temperature and humidity study
of citrus rust mite employing a constant humidity air-flow technique.
Environ. Ent. 3(5) : 813-22.
JEPPSON, L. R., J. A. MCMURTRY, D. W. MEAD, M. J. LESSER, AND H. G.
JOHNSON. 1975. Toxicity of citrus pesticides to some predaceous
phytoseiid mites. J. Econ. Ent. 68: 707-10.
KNAPP, J. L. 1983. Florida citrus spray guide. Florida Coop. Ext. Serv.
Univ. Florida Circ. 393-H. 8 p.
REED, D. K., A. K. BURDITT, JR., AND C. R. CRITTENDEN. 1964. Laboratory
methods for rearing rust mites (Phyllocoptruta oleivora and Aculus
pelekassi) on citrus. J. Econ. Ent. 57(1): 130-3.
SIMANTON, W. A., AND K. TRAMMEL. 1966. Recommended specifications for
citrus spray oils in Florida. Proc. Florida State Hort. Soc. 79: 26-30.
- -^-^ --- --- a-- -
ESTIMATING RATES OF ARTHROPOD PREDATION ON
VELVETBEAN CATERPILLAR LARVAE IN SOYBEANS
M. K. ELVIN1, J. L. STIMAC, AND W. H. WHITCOMB
Dept. of Entomology and Nematology
University of Florida
Gainesville, FL 32611 USA
ABSTRACT
A method is presented for partitioning estimates of total larval mortality
into that attributable to arthropod predators and that attributable to other
factors. Mark-release-recapture and direct observation techniques are used
in combination to derive estimates of mortality inflicted by all predators and
to identify the predator species responsible for the mortalities on small and
medium larvae. Daily rates of arthropod predation on velvetbean caterpillar,
Anticarsia gemmatalis Hiibner, larvae ranged from 0-25% per day for small
larvae and 0-73% per day for medium larvae in 2 soybean fields. Nabids
consumed the greatest proportion of smdll and medium larvae in one field,
whereas imported fire ants, Solenopsis invicta Buren, were the major con-
sumer of small and medium larvae in the other field. Results demonstrate
that the total mortality inflicted by all predators can be the same for predator
complexes with different species composition.
'Present address: Dept. of Entomology, University of Arkansas, Fayetteville, AR 72701
USA.
319
Florida Entomologist 66(3)
RESUME
Se present un m6todo de seccionar las estimaciones de la mortalidad
total de las larvas a la porci6n que se atribuye a los predadores y la que se
atribuye a otros factors. T6cnicas de marcarliberar-recapturar y de ob-
servar se usan en combinaci6n para derivar las estimaciones de la mortalidad
inflijida por todos predadores y para identificar la especie de predador re-
sponsable para las mortalidades de las larvas pequefias y de medio tamafio.
Rangos diaries de la predaci6n de artr6podos sobre larvas de Anticarisia
germmatalis Hubner variaban desde 0-25% por dia para las larvas pequefias,
0-73% por dia para las larvas de medio tamafio en 2 campos de frijol de
soya. En un campo, los nabidos consumieron la porci6n mas grande de las
larvas pequefias y de medio tamafio, mientras que en otro campo Solenopsis
invicta Buren fue el consumidor major de las larvas pequefias y las de medio
tamafio. Los resultados demonstraron que la mortalidad inflijida por todos
los predadores puede ser equal para los complejos con diferentes com-
posiciones de species.
Management programs for arthropod pests of row crops require the
ability to forecast when pest populations will reach economically injurious
levels. Since population levels are influenced by survival of progeny, predic-
tion of pest population dynamics presupposes that rates of mortality in-
flicted upon pest species may be predicted. Therefore, quantitative estimates
of pest mortality inflicted by indigenous natural enemies are needed. In this
paper we present methods for quantitatively estimating rates of stage-
specific pest mortalities due to predation by indigenous species of arthropods.
Numerous attempts have been made to quantify arthropod predation on
insect pests of row crops (Whitcomb 1981). These attempts have resulted in
2 types of information: mortality estimates for specific life stages of prey
populations and identification of the predator species known to attack a given
prey species. Mortality estimates of various prey species have been deter-
mined by using either a release-recapture technique (Luna 1979, and
McDaniel and Sterling 1982), or manipulation of prey and predator popula-
tions within field cages (Lingren et al. 1968, van den Bosch et al. 1969,
Irwin et al. 1974, and Richman et al. 1980). The arthropod predators at-
tacking a prey species have been identified by using either direct observation
(Whitcomb 1967), mark-release-capture utilizing radioisotopes as markers
(Buschman et al. 1977; McDaniel and Sterling 1979, 1982; and McCarty et al.
1980), or serology (Rothschild 1966, Greenstone 1977, Ragsdale et al. 1981,
and Gardner et al. 1981).
Mortality estimates and identification of predaceous arthropod species
determined in the aforementioned studies have limited utility in the con-
struction of dynamic models of predation because most of these studies pro-
vide only 1 of the 2 types of information needed. The construction of dynamic
models of predation requires both a description of the patterns of predator-
induced mortality and the mortality attributable to individual species under
field conditions (Stimac and O'Neil 1983). In a preliminary attempt to ob-
tain the information required to discern patterns of arthropod predation on
small and medium velvetbean caterpillar (Anticarsia gemmatalis Hiibner;
Lepidoptera: Noctuidae) larvae in soybeans, we used a method that involved
simultaneous use of mark-release-recapture of prey and direct observation
techniques. The mark-release-recapture of prey technique was used to esti-
320
September, 1983
Elvin et al.: Predation of VBC in Soybeans 321
mate the total prey mortality and the mortality inflicted by factors other
than predators. An estimate of the mortality inflicted by all predators can
be obtained by subtracting non-predator-induced mortality from total mor-
tality. By direct observation, the predator-induced mortality estimate can
then be partitioned into the mortality inflicted by each predator species. This
2-step partitioning of the total prey mortality results in a quantitative esti-
mate of the rate of arthropod predation for each predator species.
MATERIALS AND METHODS
The method for estimating rates of arthropod predation was used in 2
soybean fields, a 1-ha plot at Green Acres Agronomy Farm near Gainesville,
Florida (Green Acres site), and a 4-ha commercial field west of Archer,
Florida (Archer site). Two fields were used so that we could compare rates
of predation and predator complexes at each site.
Experiments were conducted from 30 June-22 September 1980 at Green
Acres using 'Bragg' variety soybeans planted with a 76 cm row spacing. At
the Archer site, experiments were conducted from 2 July-27 August 1980 on
'Hardee' variety soybeans planted with a 91 cm row spacing. The Archer site
was treated with methomyl on 16 August to suppress populations of velvet-
bean caterpillar (VBC) larvae. On each day the experiments were conducted,
the phenological stage of the plants was determined using the criteria of
Fehr et al. (1971).
Relative densities of predator populations within each field were esti-
mated from shake cloth samples. A sample was taken by beating the foliage
in 91 cm of 1 row onto a shake cloth spread on the ground between the rows.
Dislodged predators were either identified in the field or collected and re-
turned to the laboratory for identification. The number in each taxon was
recorded. Shake cloth samples were taken each week at 5 randomly chosen
locations within the Green Acres field. At the Archer site, 8 shake cloth
samples were taken each week using systematic sampling with a random
start (Cochran 1963).
MARK-RELEASE-RECAPTURE
Small (first and second instar; length < 12.5 mm) and medium (third
and fourth instar; length 12.5-25 mm) larvae released at the Green Acres
site were marked with radioactive phosphorus (32P) by allowing the larvae
to feed 36-48 h on 200 g of artificial diet (Greene et al. 1976) containing 2.5
ml of 5 pCi/ml 32P solution. Small and medium larvae released at the Archer
site were marked by allowing the larvae to feed 36-48 h on 200 g of artificial
diet (Greene et al. 1976) to which had been added 0.5 g of Calco Oil RedR
dye. Radioactive phosphorus could not be used at the Archer site because the
Archer site did not meet federal standards for the introduction of radio-
active materials into the environment.
To estimate total mortality, releases of each size (small and medium)
larvae were made at 2 randomly chosen sites within each field each week.
Prior to a release, the foliage at a release site was shaken to remove as many
indigenous larvae as possible. A release consisted of 20 marked larvae of 1
size placed on the foliage in 91 cm of row. Release sites were sampled 24 h
after a release using a modified clam trap (Luna et al. 1982).
Florida Entomologist 66 (3)
In each field, releases were also made into a field cage in an attempt to
estimate mortality induced by factors other than predators (background
mortality). The field cages were quonset type (3.05 m X 3.05 m X 2.26 m;
described in Collins 1980) and designed to exclude as many predators as
possible. Two days prior to a release, the foliage within a cage was vacuumed
with a D-Vac sampler. One release, consisting of 20 marked larvae placed
onto the foliage in 91 cm of row, was made within a cage each week for each
stage. The sites within a cage were sampled 24 h after a release by harvest-
ing plants and returning them to the laboratory where the foliage was care-
fully searched for larvae.
The presence of the 32P marker was determined by the use of a gas flow
windowless proportional counter. The presence of the dye was determined
by examining the fat body and posternal gland of the larvae under 10 X
magnification.
Losses due to dispersal out of the release site were considered negligible
because Luna (1979) found that small VBC larvae do not readily disperse
out of a 91 cm row area in 24 h. Medium larvae were assumed to disperse
similarly to small larvae.
DIRECT OBSERVATION
Direct observation studies, modified from that proposed by Whitcomb
(1967), were conducted concurrently with the mark-release-recapture study.
Observations were made at 20 stations set up in a circular configuration
across the field. Each station consisted of 3 leaves located in the canopy
foliage in 91 cm of row. Ten larvae were placed at each station. At 15 sta-
tions, only 1 size class was placed on the foliage (10 stations had small
larvae, and 5 stations had medium larvae). At each of the remaining sta-
tions, 5 small and 5 medium larvae were placed on the foliage. An observer
moved from station to station and recorded any predation events observed.
A predation event was defined as the successful capture of 1 prey item by
an arthropod predator(s). Before leaving a station, the observer would re-
place any missing larvae. At Green Acres, a continuous sequence of observa-
tions of the 20 stations were made from 6 am-6 pm and 7 pm-9 pm. The
stations were also checked once at midnight and at 5 am on the following
day. At the Archer site, a continuous sequence of observations were made
from 6 am-6pm.
RESULTS AND DISCUSSION
During the season at the Green Acres site, the mean daily total mor-
tality of larvae released in the field varied from 55-100% for small larvae
and from 52-83% for medium larvae (Fig. 1). The mortality of larvae re-
leased within the field cage (background mortality), representing the mor-
tality induced by factors other than predators, varied from 45-100% for
small larvae and from 25-95% for medium larvae (Fig. 1). On some sample
dates, the daily background mortality was as great or greater than the total
mortality. This was due, in part, to the inability of the field cage to exclude
predators that spent a majority of the time on the soil surface ('ground'
predators). Therefore, the daily background mortaility may have been in-
flated due to the mortality inflicted by ground predators that were not ex-
September, 1983
322
Elvin et al.: Predation of VBC in Soybeans
100 Total
S80
S 40-
I
20-
100 Background
80
40-
323
Medium
DIfOHbHIIO
40 Predator-induced
20
30 7 14 21 28 4 11 18 25 i A 15 22
JLY AUG SEP
Predator-iiduced
40-
20
30 7 14 21 28 4 11 18 25 1 8 15 22
JLY AUG SEP
Fig. 1. Estimates of mean daily mortality of velvetbean caterpillar larvae
released in the Green Acres field, 1980. R1 and R3 refer to phenological
stages of the soybeans as described by Fehr et al. (1971).
eluded by the field cage. During 8-22 September, the soybeans at Green Acres
suffered nearly complete defoliation as a result of a large population of
noctuid larvae (VBC; soybean looper, Pseudoplusia includes Walker; and
corn earworm, Heliothis zea (Boddie)). The daily mortality of larvae from
8-22 September (Fig. 1), therefore, may have been due to resource deple-
tion rather than predation.
At the Archer site, the mean daily estimates of total mortality of larvae
released in the field ranged from 48-90% for small larvae and 25-73% for
medium larvae (Fig. 2). Variability of total larval mortality estimates may
have been influenced by the presence or absence of imported fire ants,
Solenopsis invicta Buren, at the site where larvae were released. No effort
was made to avoid placing a release site n6ar a fire ant nest.
As in the Green Acres site, estimates of daily background mortality at
the Archer site, on some sample dates, were greater than estimates of mor-
tality where predators were not excluded (Fig. 2). Fire ants were the main
cause of high daily background mortality. The presence of ants within the
cage resulted from the inability of the cage to exclude fire ants and the
impracticality of searching the field for an ant-free location for cage place-
ment. To control fire ants, AmdroR bait was applied to the soil surface of
100, Total
Small
R3 Defolaltori
4
Background
I -
Florida Entomologist 66 (3)
Small
100-Total
RI Spray R3
4* 4
Background
September, 1983
Medium
Ri Spray R3
$$ *
401 Predator-induced
- 20-
Predator-induced
S401
S40
a I
a \
I-Z.
2 9 16 23 30 6 13 20 27
JLY AUG
2 9 16 23 30 6 13 20 27
JLY AUG
Fig. 2. Estimates of mean daily mortality of velvetbean caterpillar larvae
released in the Archer field, 1980. R1 and R3 refer to phenological stages of
the soybean as described by Fehr et al. (1971).
the control cage and around the outside perimeter of the cage, in an un-
successful attempt to exclude fire ants from the cage.
Mortality inflicted by all predators can be estimated by the difference
between total mortality and background mortality assuming that none of
the background mortality was due to predation. Clearly, this assumption is
violated partially when the background mortality estimates determined in
this study are used to calculate predator-induced mortality. However, the
violation of this assumption does not negate the validity of partitioning esti-
mates of total mortality using background mortality estimates. Instead, the
estimates of predator-induced mortality determined using the inflated esti-
Total
\
324
100
20-
100
80-
o
2 60-
4
g 40-
a
I / __ ~_
, . . . .
Elvin et al.: Predation of VBC in Soybeans 325
mates of background mortality are conservative estimates of the mortality
inflicted by a predator complex (DeBach et al. 1949).
To calculate the predator-induced mortality, background mortality was
subtracted from total mortality. For those dates on which background mor-
tality exceeded total mortality, estimates of predator-induced mortality were
set equal to zero. At the Green Acres site, estimates of daily mortality in-
flicted by all predators ranged from 0-25% for small larvae and 0-37% for
medium larvae (Fig. 1). The range of predator-induced mortality estimates
for larvae released at the Archer site was 0-20% per day for small larvae
and 0-73% per day for medium larvae (Fig. 2).
The direct observation technique allows the predator-induced mortality
estimates to be partitioned according to predator species. This requires esti-
mates of the mean proportion of prey consumed by a predator species on the
day of measurement. In this study we assumed that estimates made on one
day of the week reflected patterns of predation for each day of the week.
This assumption appears reasonable because densities of predators did not
vary dramatically from week to week (predator densities discussed later in
text). Results from direct observation were used to identify those species
that could be considered as the major contributors to predation mortality.
Results from direct observation are expressed as the ratio of the total
number of times a given predator species was observed consuming a prey
over the entire season to the total number of predation events observed over
the entire season. A total of 70 predation events, 36 for small larvae and 34
for medium larvae, was observed in the Green Acres field. The nabid,
Tropiconabis capsiformis (Germar), consumed the greatest proportion of
small larvae (28%, Table 1). Another nabid. Nabis roseipennis Reuter, was
observed to consume the greatest proportion of medium larvae (18%, Table
1). These results are consistent with the work of others who also found that
nabids consume relatively large proportions of immature stages of noctuid
pests of soybeans (Buschman et al. 1977, Richman et al. 1980, and Gardner
et al. 1981). In the Archer field, 53 predation events were observed, 12 for
small larvae and 41 for medium larvae. The imported fire ant consumed the
greatest proportion of small (75%, Table 2) and medium larvae (87.5%,
Table 2).
Comparisons of the predation by each species (Tables 1 and 2) suggest
that imported fire ants, when present, dominated the predation on VBC
larvae. Their dominance, however, did not appear to extend to the species
composition of the predator complex because the densities of the species
comprising the complex within the 2 fields were similar (Table 3). To deter-
mine if the addition of imported fire ants to the complex resulted in differ-
ences in the rates of mortality, comparisons of the estimates of daily total
mortality were made using a 2-way ANOVA procedure with location and
weeks as factors. No statistically significant differences (P = 0.10) in the
rates of daily total mortality for small larvae and medium larvae could be
detected between the 2 fields or among weeks. Patterns of daily background
mortality were similar also (Fig. 1 and 2). This suggests that imported fire
ants inhibit the foraging activity of other predator species, yet compensate
for this inhibition by inflicting comparable amounts of mortality on a pest
population. Imported fire ants also have been shown to play a similar role in
the dynamics of Heliothis virescens (F.) in Texas cotton (Sterling et al.
Florida Entomologist 66 (3)
TABLE 1.
THE SEASONAL AVERAGE PROPORTION OF LARVAE PREYED UPON BY A
PREDATOR SPECIES AS DETERMINED BY DIRECT OBSERVATION IN THE
GREEN ACRES SITE. THE TOTAL NUMBER OF LARVAE PREYED UPON
WAS 70 (36 SMALL LARVAE AND 34 MEDIUM LARVAE).
Predator species
Proportion consumed
Small Larvae
Tropiconabis capsiformis (Germar)
Geocoris punctipes (Say)
Theridion albidum Banks
Misumenops spp.
Hentzia palmarum (Hentz)
Nabis roseipennis Reuter
Chiracanthium spp.
Tettigoniidae
Podisus maculiventris (Say)
Labidura ripara (Pallas)
Medium Larvae
Nabis roseipennis Reuter
Misumenops spp.
Labidura ripara (Pallas)
Peucetia viridans (Hentz)
Calleida decora (F.)
Chiracanthium spp.
Geocoris punctipes (Say)
Hentzia palmarum (Hentz)
Tettigoniidae
Tropiconabis capsiformis (Germar)
Podisus maculiventris (Say)
Tetragnathus spp.
Oxyopes salticus Hentz
Theridion albidum Banks
0.28
0.19
0.11
0.08
0.08
0.08
0.06
0.06
0.06
0.03
0.18
0.15
0.15
0.09
0.09
0.09
0.06
0.03
0.03
0.03
0.03
0.03
0.03
0.03
TABLE 2. THE SEASONAL AVERAGE PROPORTION OF LARVAE PREYED UPON BY A
PREDATOR SPECIES AS DETERMINED BY DIRECT OBSERVATION AT THE
ARCHER SITE. THE TOTAL NUMBER OF LARVAE PREYED UPON WAS
53 (12 SMALL LARVAE AND 41 MEDIUM LARVAE).
Predator species Proportion consumed
Small Larvae
Solenopsis invicta Buren 0.75
Conomyrma spp. 0.17
Hentzia spp.1 0.08
Medium Larvae
Solenopsis invicta Buren 0.88
Conomyrma spp. 0.03
Tettigoniidae 0.03
Oxyopes spp.1 0.02
Chiracanthium inclusum (Hentz) 0.02
Clubionidae1 0.02
1Spider immature-not identifiable to species.
326
September, 1983
TABLE 3.
Elvin et al.: Predation of VBC in Soybeans 327
MEAN NUMBER OF ARTHROPOD PREDATORS FOUND IN SHAKE CLOTH
SAMPLES FROM 30 JUNE-27 AUGUST AT THE GREEN ACRES AND
ARCHER SITES. MEANS AT THE GREEN ACRES SITE ARE BASED ON 5
SAMPLES PER WEEK, AND THOSE FOR THE ARCHER SITE, 8 SAMPLES
PER WEEK.
Predator categories,
Hemi- Cara-
Date Spiders Ants ptera bidae Others
30 June-2 July
Green Acres 0 0 1.0 0 0.2
Archer 3.1 0.3 4.9 0 0.3
7 July-9 July
Green Acres 0.2 0 1.6 0 0
Archer 1.0 0.6 0.9 0 0
14 July-16 July
Green Acres 0.6 0.2 0.8 0 0
Archer 0.6 0.8 1.2 0.1 0
21 July-23 July
Green Acres
Archer
0.8 0.2 0.8
- RAIN
28 July-30 July
Green Acres
Archer
4 August-6 August
Green Acres
Archer
11 August-13 August
Green Acres
Archer
18 August-20 August
Green Acres
Archer
25 August-27 August
Green Acres
Archer
'Species and/or families
from the authors.
of organisms included in these categories available upon request
1979; McDaniel and Sterling 1979, 1982; and Agnew et al. 1982). In gen-
eral terms, the results of this study and that done in Texas cotton suggest
that the mortality inflicted upon prey populations is not necessarily de-
pendent upon the presence or absence of a particular species within the
complex. The species comprising the complex may be able to compensate for
one another in terms of the mortality they inflict upon a prey population.
The generality of such an effect, however, must be the subject of further
study.
0 0.2
328 Florida Entomologist 66 (3) September, 1983
CONCLUSIONS
Quantitative estimates of the rates of arthropod predation are pre-
requistes for the construction of models of pest dynamics. The method used
in this study provided the quantitative estimates of arthropod predation
rates by partitioning total mortality estimates into predator-induced and
non-predator-induced components, and identification of the arthropod preda-
tor species that could be considered as the major species responsible for the
mortality. Partitioning of the mortality estimates and identifying the major
predator species are accomplished through the simultaneous use of mark-
release-recapture and direct observation techniques. With slight modification
of the techniques, this method could be used to estimate rates of arthropod
predation on other noctuid pests of row crops.
The seasonal mean rates of arthropod predation reported in this study
were 13.8% ( 1.7%) mortality per day for small VBC larvae and 28.5%
( 6.4%) mortality per day for medium VBC larvae. These rates should
be considered as conservative estimates of predator-induced mortality be-
cause of the inflated estimates of background mortality used in their calcula-
tion. Assuming that these rates are imposed every day during the develop-
ment of small and medium larvae (7 days, Reid 1975), only 17 individuals
of a cohort of 100 newly emerged larvae would survive to the end of the
medium stage. The number of small and medium larvae would be reduced
by 83% by the end of the fourth instar even at these rates of predation.
Comparisons of the mortality estimates, observation records, and species
composition of the predator complexes in the 2 fields suggest that the amount
of mortality inflicted upon prey populations is not necessarily dependent
upon the presence or absence of a particular predator species in the com-
plex. Although the densities and species composition of the predator com-
plexes varied, the patterns of predator-induced mortality were very similar.
However, further study is needed to determine the generality of a com-
pensatory effect demonstrated by the species comprising the predator com-
plex.
ACKNOWLEDGMENTS
This research was supported by EPA Grant No. CR-806277-02-0, "The
Development of Comprehensive, Unified, Economically and Environmentally
Sound Systems of Intergrated Pest Management For Soybean."
We wish to acknowledge Drs. H. Cromroy and B. Smittle for providing
the laboratory facilities necessary for handling radioactive materials. We
would also like to acknowledge the technical assistance provided by Dr.
R. Hemenway, Ms. L. Daniels, Mr. R. Barzel, and Mr. H. Robles. We would
like to thank Drs. D. Richman, C. Barfield, D. Habeck, and S. Kerr for re-
viewing the manuscript. University of Florida Agricultural Experiment
Station Journal Series No. 3830.
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AGNEW, C. W., W. L. STERLING, AND D. A. DEAN. 1982. Influence of cotton
nectar on red imported fire ants and other predators. Environ. Ent.
11: 629-34.
BUSCHMAN, L. L., W. H. WHITCOMB, R. C. HEMENWAY, D. L. MAYS, NGUYEN
Elvin et al.: Predation of VBC in Soybeans
Ru, N. C. LEPPLA, AND B. J. SMITTLE. 1977. Predators of velvetbean
caterpillar eggs in Florida soybeans. Environ. Ent. 6: 403-7.
COCHRAN, W. G. 1963. Sampling techniques. John Wiley and Sons, Inc.,
New York.
COLLINS, F. L. 1980. Response of a predator complex to larval density of
Anticarsia gemmatalis (velvetbean caterpillar) and Pseudoplusia
includes (soybean looper). M.S. Thesis. University of Florida,
Gainesville, Florida. 161 p.
DEBACH, P., E. J. DIETRICK, AND C. A. FLESCHNER. 1949. A new technique
for evaluating the efficiency of entomophagous insects in the field. J.
Econ. Ent. 42: 546-7.
FEHR, W. R., C. E. CAVINESS, D. T. BRUMOOD, AND J. S. PENNINGTON. 1971.
Stage of development descriptions for soybean, Glycine max (L.)
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GARDNER, W. A., M. SHEPARD, AND R. NOBLET. 1981. Precipitin test for ex-
amining predatory-prey interactions in soybean fields. Canadian
Ent. 113: 365-9.
GREENE, G. L., N. C. LEPPLA, AND W. A. DICKERSON. 1976. Velvetbean cater-
pillar: A rearing procedure and artificial medium. J. Econ. Ent. 69:
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GREENSTONE, M. H. 1977. A passive haemagglutination inhibition assay for
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IRWIN, M. E., R. W. GILL, AND D. GONZALEZ. 1974. Field-cage studies of
native egg predators of the pink bollworm in southern California
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LINGREN, P. D., R. L. RIDGWAY, AND S. L. JONES. 1968. Consumption by
several common arthropod predators of eggs and larvae of two
Heliothis species that attack cotton. Ann. Ent. Soc. America 61: 613-8.
LUNA, J. M. 1979. A tactical economic threshold model for velvetbean cater-
pillar (Anticarsia gemmatalis Hbn.) in Florida soybeans. M.S. Thesis.
University of Florida, Gainesville, Florida. 102 p.
H. M. LINKER, J. L. STIMAC, AND S. L. RUTHERFORD. 1982. Estima-
tion of absolute larval densities and calibration of relative sampling
methods for velvetbean caterpillar (Anticarsia gemmatalis Hbn.) in
soybean. Environ. Ent. 11: 497-502.
MCCARTY, M. T., M. SHEPARD, AND S. G. TURNIPSEED. 1980. Identification of
predaceous arthropod species in soybeans using autoradiography.
Environ. Ent. 9: 199-203.
MCDANIEL, S. G., AND W. L. STERLING. 1979. Predator determination and
efficiency on Heliothis virescens eggs in cotton using 32P. Environ.
Ent. 8: 1083-7.
1982. Predation of Heliothis virescens (F.) eggs on cotton in East
Texas. Environ. Ent. 11: 60-6.
RAGSDALE, D. W., A. D. LARSON, AND L. D. NEWSOM. 1981. Quantitative
assessment of the predators of Nezara viridula eggs and nymphs
within a soybean agroecosystem using an ELISA. Environ. Ent. 10:
402-5.
REID, J. C. 1975. Larval development and' consumption of soybean foliage by
the velvetbean caterpillar, Anticarsia gemmatalis Hbn. (Lepidoptera:
Noctuidae), in the laboratory. Ph.D. Dissertation. University of
Florida, Gainesville, Florida. 125 p.
RICHMAN, D. B., R. C. HEMENWAY, AND W. H. WHITCOMB. 1980. Field cage
evaluation of predators of the soybean looper, Pseudoplusia includes
(Walker) (Lepidoptera: Noctuidae). Environ. Ent. 9: 315-7.
ROTHSCHILD, G. H. L. 1966. A study of a natural population of Conomelus
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330 Florida Entomologist 66 (3) September, 1983
anceps (Germar) (Homoptera: Delphacidae) including observations
on predation using a precipitin test. J. Anim. Ecol. 35: 413-34.
STERLING, W. L., D. JONES, AND D. A. DEAN. 1979. Failure of the red
imported fire ant to reduce entomophagous insect and spider abundance
in a cotton agroecosystem. Environ. Ent. 8: 976-81.
STIMAC, J. L., AND R. J. O'NEIL. 1983. Modeling the impact of natural
enemies on insect pests of soybeans. Pages 78-87 In H. N. Pitre, ed.
Natural enemies of arthropod pests of soybeans. Southern Reg. Bull.
No. 285.
VAN DEN BOSCH, R. T., T. F. LEIGH, S. GONZALEZ, AND R. E. STINNER. 1969.
Cage studies of predators of the bollworm in cotton. J. Econ. Ent.
62: 1468-9.
WHITCOMB, W. H. 1967. Field studies on predators of second-instar boll-
worm, Heliothis zea (Boddie) (Lepidoptera: Noctuidae). J. Georgia
Ent. Soc. 2: 113-8.
.1981. The use of predators in insect control. Pages 105-123 In
D. Pimentel, ed. Pest management. Vol. II. CRC Press, West Palm
Beach, Florida.
BEHAVIORAL ECOLOGY OF MATING IN THE
CARIBBEAN FRUIT FLY, ANASTREPHA SUSPENSE
(LOEW) (DIPTERA: TEPHRITIDAE)
THEODORE BURK1
Insect Attractants, Behavior, and Basic Biology Research Laboratory,
Agric. Res. Serv., USDA, Gainesville, FL 32604 USA
and
Department of Entomology and Nematology, Univ. of Fla.
Gainesville, FL 32611 USA
ABSTRACT
Observations on the behavioral ecology of Caribbean fruit flies,
Anastrepha suspense (Loew), were made during spring and summer 1981 on
Surinam cherry (Eugenia uniflora L.) and guava (Psidium guajava L.)
host trees. General activity patterns were as follows: males and females
were active on fruit in the early morning (males feeding and courting,
females feeding and ovipositing), but moved to positions under shaded leaves
in the peripheral canopy in mid-morning as light intensity and temperature
increased. In the late afternoon males fought for single-leaf territories,
"puffed" to produce sex pheromone, and "called" with acoustic signals. Al-
most all matings took place following attraction of females to these display-
ing males. Males therefore had 2 mating modes, searching on fruit and dis-
playing under leaves. Territorial fights were an important part of the more
profitable mode. Large size and residence were correlated with success in
fights.
RESUME
Se hicieron unas observaciones sobre la ecologia de comportamiento de la
mosca caribe de la fruta, Anastrepha suspense (Loew) durante la primavera
1Current address: Department of Biology, Creighton University, Omaha, NE 68178.
Burk: Mating in Caribbean Fruit Flies
y el vcrano de 1931 robra los arbolbs hospederos, la cereza de Surinam,
Eugcnia uniflora L. y la guayaba, Psidium guajava L. Los patrons generals
de actividad fueron como en seguida: las hembras y los machos fueron
activos sobre las frutas por la mariana temprano (los machos alimentandose
y cortejando, las hembras alimentandose y ovipositando), pero movieron a
posiciones debajo la sombra de hojas en el pabell6n perif6rico en el medio
de la mafiana mientras la intensidad de la luz y la temperature se aumen-
taron. Por la tarde, los machos pelearon para territories de particulars
hojas, producieron feromona sexual, y llamaban con sefiales acusticos. Casi
todos los apareamientos tuvieron lugar despues de la atracci6n de las hembras
por estos machos demonstrativos. Asi los machos tuvieron dos modos de
aparear: buscar sobre las frutas y demonstar debajo de las hojas. Los vuelos
territoriales formaron una parte important del modo mis provechoso. El
tamano grande y la residencia se relacionaron con los 6xitos en los vuelos.
Tephritid fruit flies can be classified into 2 general types on the basis of
their host relations and mating systems. Monophagous species, often
temperate in distribution, usually mate where the female oviposits (fruit or
site of gall formation). Polyphagous species, often tropical or subtropical,
usually mate in aggregations on the foliage of a host plant, but not neces-
sarily in the immediate vicinity of the oviposition resource (Prokopy 1980,
Burk 1981). There are exceptions to these broad generalizations (Smith and
Prokopy 1980).
There have been a number of excellent field studies of the mating be-
havior of tephritids of the first type, especially of the genus Rhagoletis by
Prokopy and co-workers (reviewed in Prokopy 1980, Smith and Prokopy
1980). Most studies of the behavior of the second type have been conducted
either in the laboratory or with flies released onto field-caged host trees
(Tychsen 1977, Prokopy and Hendrichs 1979, Suzuki and Koyama 1981,
Dodson 1982). While these methods have yielded important findings, espe-
cially in terms of descriptions of behavior and quantitative data, it is de-
sirable to supplement them with observations on undisturbed wild popula-
tions, particularly in view of the high densities maintained in some of the
controlled situations.
In this paper I report a study of a wild population of the Caribbean fruit
fly, Anastrepha suspense (Loew) (="caribfly"), in South Florida. Like the
medfly, the caribfly is extremely polyphagous (records from over 100 hosts,
Swanson and Baranowski 1972). This study was made in conjunction with a
field-cage study of released caribflies by J. Hendrichs (unpubl. data).
There have been several previous reports on aspects of caribfly mating
behavior. Nation (1972) described courtship and mating behavior and re-
ported the production by males of an airborne sex pheromone. Perdomo
(1974) described mating interactions in a wild population and showed that
under field conditions male sex pheromone was equally attractive to female
and male caribflies. Webb et al. (1976) described calling, precopulatory, and
aggressive sounds produced by rapid wing vibrations of males. Dodson
(1982), in a field-cage study, emphasized that displaying males tended to
aggregate (also noted by Perdomo), and described aggressive interactions
between males over "territories" on the underside of leaves in host trees.
My observations were designed to obtain an integrated picture of caribfly
331
332 Florida Entomologist 66 (3) September, 1983
mating behavior and to examine aspects of sexual selection (male-male
competition and female mate choice) in an undisturbed caribfly population.
METHODS
Observations were made on a naturally-established caribfly population
in Homestead, FL. This was the area in which caribflies were observed by
Perdomo (1974). In May and June 1981, flies were observed on a 40-m long,
3-m high hedge of 22 Surinam cherry trees (Eugenia uniflora L.) on the
grounds of the University of Florida, Institute of Food and Agricultural
Sciences (IFAS), Agricultural Research and Education Center (AREC).
In July-October 1981, observations were made in a grove of guava trees
(Psidium guajava L.) on the Sandoz Farm adjacent to the IFAS, AREC.
This site was ca 400 m from the Surinam cherry site. Surinam cherries
and guavas are the preferred hosts of caribflies in the late spring and late
summer, respectively. Studies on various tephritids show them to be active
dispersers (Bateman 1972); caribflies studied on the 2 hosts were almost
certainly part of the same genetic population.
The main observation method was to walk a transect line each hour
from 0700 to 1900, Eastern Daylight Time (EDT), recording the sex, loca-
tion and behavior of all caribflies seen. The route entailed walking on each
side of the Surinam cherry hedge in the spring, and on each side of one
60-m row of 10 guava trees in the summer. These hourly census counts
usually took about 30 min; the remainder of an hour was spent observing
flies using the "focal animal" technique (Altman 1974). A fly was selected at
random and observed for 5 min, during which its behavior and social inter-
actions were noted. Then another fly was observed for 5 min, and so on. At
the beginning of each count the temperature, approximate wind speed, and
light intensity were noted. Temperature was recorded with a Fisher Sci-
entific (Pittsburg, PA) hand-held digital thermometer, light intensity with
a General Electric Model 214@ triple range light meter. Measurements were
made under a shaded leaf in the peripheral canopy of the host trees. During
the period from 26 May to 7 October, 214 hourly census counts were made.
Counts were not made during showers, when flies were inactive. Also,
journey time to the study area at beginning and end of observation weeks
meant that certain hours included observations less often than others. Actual
numbers of census counts are given in Table 1. In the presentation of the
results, counts are given according to EDT. Since sunrise and sunset varied
over this period, this introduces slight variability into the results. However,
as the results will show, clear patterns of activity emerged nonetheless.
OBSERVATIONS
CENSUS COUNTS AND GENERAL ACTIVITY PATTERNS. No obvious differences
were noted in the daily abundance or activity patterns of caribflies on
Surinam cherries and guavas, so data from the 2 hosts were combined.
The average number of caribflies observed in hourly census counts is
shown in Fig. 1. The number of flies observed increased during the morning
hours and remained relatively steady from 1000 on. Usually a slight excess
of females was observed, but equal numbers of males and females at 1700 h
suggests that this was due to greater visibility of females than males at
Burk: Mating in Caribbean Fruit Flies
CARIBFLY CENSUS COUNTS
A
7 8 9 10 11 12 13
TIME (EDT)
14 15 16 17 18 19
Fig. 1. Average numbers of caribflies observed in hourly
Surinam cherry and guava host trees, Homestead, FL 1981.
censuses on
times other than the late afternoon. The peak in male numbers at 1700 h
corresponds with peaks in male sexual activities, described below.
The main factors affecting caribfly activity seemed to be light intensity
and temperature. Both male and female flies were active on fruit in the
early morning when light intensity and temperature were low (Fig. 2a,
Table 1). As the sun rose righer, flies moved to positions under leaves in the
peripheral canopy (Fig. 2b). Subsequently, it was extremely unusual to see
a caribfly in direct sunlight or even underneath a leaf whose top surface was
exposed to direct sunlight. The shift from fruit to leaves was usually almost
complete by 1000 h, but on cool or cloudy mornings was delayed until 1100 h
or 1200 h. During periods of high winds or showers, flies remained inactive
under leaves. Female flies often returned to fruit in the very late afternoon
(1900 h).
ACTIVITIES ON HOST FRUIT AND UNDER LEAVES. The locations of male and
female caribflies were similar and shifted in a coordinated manner (Fig.
2a,b). On fruit, female flies were usually either feeding or ovipositing,
while male flies were either feeding or following and courting females.
Ovipositing females moved over the surface of host fruits, "licking" the
surface with their proboscis and probing with their ovipositor. On a suitable
host, females oviposited while adopting a characteristic posture with the
ovipositor tip puncturing the fruit. After ovipositing, females dragged their
ovipositors on the fruit surface, presumably depositing a pheromone to deter
subsequent ovipositions (Prokopy et al. 1977). Females moving on the fruit
surface prior to ovipositing were probably testing for the presence of
333
334
100-
I
10
100r
Florida Entomologist 66 (3)
A) CARIBFLIES ON FRUIT
B)CARIBFLIES UNDER LEAVES
I I I l l I I I I I
7 8 9 10 11 12 13 14 15 16
17 18 19
TIME (EDT)
Fig. 2. Location of observed caribflies in host trees. (A) Percentage of
observed flies that were on host fruit. (B) Percentage of observed flies that
were under leaves in host trees.
I
September, 1983
Burk: Mating in Caribbean Fruit Flies 335
oviposition-deterring pheromone, as well as determining suitable places to
insert the ovipositor. Relatively few females were actually seen ovipositing
(ovipositor inserted into fruit), but many were seen probing or dragging.
All suitable fruit were infested with caribfly larvae. There was an early
morning peak in oviposition activity, a gradual decline throughout the middle
of the day, and a slight increase very late in the afternoon (Fig. 3).
Oviposition on guavas was almost always seen on small, hard, dark green
fruit (approximately the size of a golf ball). Fights between females,
probably over access to oviposition sites, were occasionally observed on such
fruit. Feeding flies were found on mature, pale-green-to-yellow, baseball-
sized fruit that often had large cracks and splits oozing fluid. Feeding flies
were often seen in groups (up to 10 flies) on fruit with splits. Such feeding
groups often included male flies; other males courted both feeding and
ovipositing females.
Males courting females on fruit were very active, orienting to and
following females and constantly waving their wings. These wing move-
ments consisted of alternate presentations of the ventral surface of the
wings, costal edge up (detailed description of wing movements in Nation
1972). Courting males on fruit were very common and accounted for ap-
proximately half the males seen on fruit. Only 2 of these courtships led to
attempted mounts, and no copulations resulted. The only mating pair ob-
served on fruit was in the late afternoon, and the premating behavior of
this pair was not observed. Caribflies rarely move after coupling, so courtship
in this case probably took place on the fruit.
After about 1000 h, most caribflies were observed on the underside of
leaves in the peripheral canopy, about 1-3 m above the ground. In most cases,
only one fly was seen on a leaf. Females usually remained quiet under leaves
throughout the afternoon. Males were generally inactive through the middle
of the day, but after 1500 h they became increasingly active. Males moved
from leaf to leaf, interacted aggressively, released sex pheromone and pro-
15OVIPOSITION
0 10-
0
O+
0+
ZI 5
O-
0
7 8 9 10 11 12 13 14 15 16 17 18 19
TIME (EDT)
Fig. 3. Temporal occurrence of female oviposition behavior. Number of
females seen with ovipositors inserted in fruit = 74.
Florida Entomologist 66 (3)
TABLE 1. NUMBER OF CENSUS COUNTS, TEMPERATURE AND LIGHT INTENSITY.
26 MAY-7 OCTOBER 1981. HOMESTEAD, FL.1
Mean light
Number of Temperature (oC) intensity
Time census counts Mean Range Lux (10-3)
0700 12 24.2 23-26 1.20
0800 19 26.6 26-28 4.65
0900 18 28.5 26-31 5.57
1000 16 29.7 27-31 11.32
1100 16 30.8 30-33 14.80
1200 12 30.8 29-34 11.71
1300 14 30.6 26-35 19.87
1400 16 31.8 27-33 13.58
1500 19 31.4 30-34 11.14
1600 18 30.3 29-35 9.90
1700 23 31.8 26-33 7.39
1800 21 30.0 28-32 7.18
1900 10 30.3 29-32 4.09
'Temperature and light intensity measured once per census under a shaded leaf in peri-
pheral canopy.
duced calling sounds. Most mating pairs were observed after 1600 h. These
mating-related behaviors are described more fully below.
MALE TERRITORIALITY AND SEXUAL DISPLAYS. Dodson (1978) first described
aggressive interactions between male caribflies in the context of extended
residence on a single leaf by an individual male, where the male performed
various sexual displays. My observations confirmed that territoriality is an
important part of the reproductive behavior of male caribflies.
Caribflies resting under a leaf adopt a characteristic "resting" posture,
in which the wings are folded over one another on the fly's back. Most fe-
males seen under leaves adopted this posture, while the majority of males
under leaves were in an "alert" posture, raised further off the leaf surface
on their legs with wings held at an acute angle to the line of the body axis.
As the afternoon progressed, males were increasingly observed in a char-
acteristic location under leaves; at the leaf base, facing the leaf stem, often
standing on the main leaf vein. Alert males did not perform sexual displays,
but did orient to and follow the movements of caribflies and other insects,
and reacted to other caribflies landing on their leaf territory. As males be-
came more active, they began to "wing-wave" as previously described for
courting males on fruit, often "sidling" from side-to-side on the leaf.
Eventually, usually by 1600 h, males began vigorous sexual displays. These
started with sex pheromone release, indicating by "puffing" (expansion of
the pleural regions of the abdomen), version of moist rectal tissue, and
repeated touching of the leaf surface by the abdomen tip. At this time,
males held their wings stiffly perpendicular to the body axis, costal edge up.
Eventually most puffing males began acoustic calling by production by short
bursts of rapid wing vibration of stereotyped acoustic signals (Webb et al.
1976). Calling males pivoted between bursts of wing vibration and continued
to mark the substrate with pheromone. All calling males were simultaneously
puffing, but not all puffing males called. Male puffing and calling reached a
peak at 1700-1800 but occasionally occurred earlier in the day (Fig. 4b).
336
September, 1983
Burk: Mating in Caribbean Fruit Flies 337
Displaying males were often interrupted by other males flying onto the
leaf territory. Such "intruders" either flew in from adjacent leaf territories
or from undetermined locations farther away. Reactions of resident males
to intruders have been described by Dodson (1982). I found that intruders
usually approached residents, rather than vice versa. Intruders approached
by sidling and wing-waving. Residents turned to face intruders. Residents
adopted a characteristic defense posture, the body tightly pressed to the
substrate and stretched to full length, the wings tightly pressed against the
side of the body, and the proboscis extended forward. The general shape was
of an arrowhead, with the proboscis at the point facing the intruder. Resi-
dents oriented to the movements of intruders, but usually stayed at the base
of the leaf. If intruders did not fly away when threatened by the adoption
of the defense posture by residents, but approached and contacted residents,
a fight ensued. (Most male-male encounters on leaves did result in fights.)
The 2 males butted heads and pushed against one another, with one or both
males producing long bursts of rapid wing vibration (aggressive songs).
Unlike calling songs, these aggressive songs were clearly audible and some
fights were first detected by hearing them. After a few seconds and perhaps
several pushing and buzzing bouts, one male flew away. Rarely both males
fell off the leaf or flew away. Sometimes both fell off and one returned.
Rarely did 2 males remain on a single leaf longer than a minute or 2.
Dodson (1982) was unable to distinguish clearly between residents and
intruders, but felt that residents may have had an edge in contests. Provided
I saw the 2 males before actual physical contact, I was able to distinguish
residents and intruders. Table 2 shows that Dodson's suspicion was correct;
residents are significantly more successful in territorial fights than intruders.
I observed 89 male-male encounters under leaves; in 81 cases a resident and
intruder were identified. In these 81, the intruder alone flew away 80% of the
time, the resident alone 15%, and both males 5%. It was not possible in these
field observations to evaluate the relative sizes of interacting males, but in
controlled situations (Burk unpublished observation, Hendrichs unpublished
data) it has been determined that large males have an advantage over small
ones. In these situations fight outcomes are determined by the interaction of
the size and residence effects. That is, large residents win most often, fol-
lowed by small residents, large intruders, and small intruders.
In 4 encounters residents and intruders were not identified: all ended
with one fly leaving the leaf. Four encounters involved adjacent residents
each standing on his own leaf; in all cases one resident flew away.
Eight encounters were seen in the morning on fruit. These were marked
by much lower intensity of aggression: in 4 cases both males remained on
the fruit.
The timing of male aggressive encounters is shown in Fig. 4a. The peak
in male fights occurred at 1700-1800 h. This corresponded with the peak in
sexual display (Fig. 4b), showing the coincidence between male aggressive
and sexual behavior in caribflies.
MATINGS. Although males fight proximately for possession of single-leaf
territories, ultimately these fights settle access to receptive females. Of 26
matings observed during this study, 24 took place under leaves in the loca-
tions where males fight and display. One other took place on the upper sur-
face of a leaf and the remaining one on a Surinam cherry. The temporal
Florida Entomologist 66 (3)
A) MALE FIGHTS
September, 1983
338
4C
z
S3C
u
z
U
UJ
20
X 10
ZI
8
S6
)-
04
Q 4
UJ
-i
2
Z'
IX
C) MATINGS
B) MALE DISPLAY
15-
10-
%n
Z
1-
1-
HnnR
I I 1 I I I T I 1 1 1 I
7 8 9 10 11 12 13 14 15 16 17 18 19
TIME (EDT)
'
| |
Burk: Mating in Caribbean Fruit Flies
TABLE 2. Anastrepha suspense MALE-MALE ENCOUNTERS ON SURINAM
CHERRY AND GUAVA HOST TREES. 26 MAY-7 OCTOBER 1981. HOME-
STEAD, FL.
Location Type of encounter Outcomel N
On fruit Intruder approaches Both stay 4
I stays 1
R stays 2
I, R ?, one stays 1
Under leaves Adjacent residents One goes 4
Intruder approaches R stays 65
I stays 12
Both go 4
I, R ?, one stays 4
Total 97
1I = Intruder; R = resident.
occurrence of matings coincided with male territorial fighting and sexual
displaying: 22 of 26 matings (85%) occurred between 1700 and 1900 h.
It was not possible in this study to analyze in detail the mechanisms of
female choice of mates, other than the preference for displaying, territory-
holding males. Only 2 matings were observed from the outset. In both cases,
a female landed on the underside of a leaf where a male was puffing and
calling. The male turned to face the female, and she approached waving her
wings. When the 2 made head-to-head contact, the male flew over the female,
turned around, and mounted. After mounting, the male produced a long con-
tinuous rapid wing vibration (precopulatory song). The female raised her
ovipositor, allowing male intromission. Mate choice in these 2 matings was
clearly performed by the female. These 2 matings lasted 30 5 min. Court-
ship and mating behavior have been more completely described by Nation
(1972), Perdomo (1974), and Dodson (1978).
Mating pairs often occurred spatially and temporally clumped to a
greater degree than one would expect by chance. For example, on both 6 and
7 October, mating pairs were observed simultaneously within 30 cm of each
other.
(E) PREDATORS. A large variety of potential predators was present in the
Surinam cherry and guava trees: spiders, ants, wasps, dragonflies, scorpion-
flies, anoles, and birds, among others. The most significant predation was
probably by spiders. One male caribfly was seen in the web of an unidentified
spider. Two males and 3 females were seen in the grasp of small green
salticid spiders, Lyssomanes viridis (Waeckenaer). On 8 June, I watched as
one of these spiders stalked calling males. It oriented to the movements of
males and jumped from leaf to leaf pursuing them. On that occasion, no
caribflies were captured. Hendrichs (unpubl. data) found that significant
Fig. 4. Caribfly sexual behavior. (A) Timing of male aggressive inter-
actions. (B) Average number of puffing and calling males/census count.
(C) Timing of matings. N mating pairs observed = 26.
339
340 Florida Entomologist 66 (3) September, 1983
numbers of his released caribflies were caught by spiders once he stopped
removing spiders and spider webs from his field-caged guava tree.
DISCUSSION
MATING BEHAVIOR ON FRUIT
This study reports for the first time the probable existence of dual mating
modes by male caribflies. Such alternatives have been discovered in a number
of animals (Cade 1980) including flies (Maier and Waldbauer 1979, Prokopy
and Hendrichs 1979, Smith and Prokopy 1980). In several monophagous
species of Rhagoletis, alternative male patterns are related to host phenol-
ogy: early-season matings take place on host vegetation, while late-season
matings result from males courting ovipositing females on host fruit. A
"search-on-fruit, display-on-leaves" dichotomy exists in the polyphagous
Mediterranean fruit fly, although in this case males signal daily from leaves
in the late morning to early afternoon, and court on fruit before and after
this period. The caribfly pattern resembles that of the medfly, but males
court on fruit only in the morning and signal from leaves in the late after-
noon.
In cases where dual male mating patterns occur, it is useful to consider
whether the alternatives are equally successful. Alternatives can be main-
tained in a population because they are performed under different circum-
stances; because of frequency-dependent selection leading to equal lifetime
reproductive success; or because secondary modes are performed by males
that are excluded from adopting the primary mode, "making the best of a
bad job" (Dawkins 1980).
In the apple maggot, Rhagoletis pomonella (Walsh), mating attempts by
males were as successful on leaves (39/68 = 57%) as on fruit (62/111 =
56%) (Smith and Prokopy 1980). Most mating attempts on leaves began
with males facing females and were unresisted; however, most mating at-
tempts on fruit began with males behind ovipositing females that resisted at
first. Matings on leaves are probably mostly with virgin females, matings on
fruit mostly with previously-mated females. Sperm displacement favoring
fertiliation of eggs by the last male to mate has been shown in the apple
maggot (Myers et al. 1976).
In the caribfly and medfly, however, matings on fruit seem to be much
rarer than matings under leaves. Prokopy and Hendrichs (1979) observed
39 medfly matings under leaves (50% of encounters resulted in attempted
copulations, of which 74% succeeded). Only 7 matings took place on fruit
(24% of encounters resulted in attempted copulations, of which only 23%
succeeded). In my caribfly observations, only 2 attempted copulations were
seen on fruit (neither succeeded), while one mating pair was observed on
fruit. Both medfly and caribfly females on fruit seem to be able to success-
fully resist the advances of courting males. In this study, 2 of 2 attempted
copulations under leaves succeeded, and 25 mating pairs were observed under
or on leaves. Calling males under leaves are perhaps subject to greater
predation from spiders than courting males on fruit, but even so it is very
likely that displaying is the far more profitable mating mode in these 2
species.
The less successful mode, searching on fruit, might be a strategy pursued
Burk: Mating in Caribbean Fruit Flies
by territorial males at times when they are not displaying, or it could be
pursued by males that are excluded by male-male competition from holding
display territories. I am unable to distinguish between these 2 possibilities,
but Hendrichs (unpubl. data) has some evidence which suggests that males
that were less successful in territorial fights were more often found courting
on fruit.
Prokopy and Hendrichs (1979) found that virgin female medflies as well
as mated ones engaged in oviposition activities on fruit. I observed significant
amounts of feeding on fruit by female caribflies, probably including virgins.
Both of these observations suggest that matings on fruit could involve virgin
females as well as already-mated ones. Since unresponsive medfly and
caribfly females usually seem able to resist forced matings, the presence of
virgin females on fruit could help to explain male courtship on fruit in the
medfly and caribfly as a secondary mating strategy.
MATING BEHAVIOR UNDER LEAVES AND IMPORTANCE OF MALE COMPETITION
Prokopy (1980) and Burk (1981) have suggested that matings away
from fruit in multivoltine polyphagous tephritids are due to greater vari-
ability and less predictability of host fruit resources. This would result in
fewer encounters with females by males defending territories on fruit. The
presence of oviposition-deterring pheromones on fruit already visited by
females may also reduce the amount of time females are present on fruit and
the likelihood of their adopting oviposition postures which make them vul-
nerable to males. These factors may have been important in the evolution of
mating on vegetation in species such as the medfly and caribfly.
Males displaying in vegetation maximize the number of females attracted
by displaying almost exclusively in host trees. I occasionally saw calling
males in nonhosts, but these were always adjacent to host trees with numbers
of caribflies present. In addition to relying on visual responses of females to
host trees, male caribflies attract females through the production of at-
tractant pheromones and acoustic signals.
In field-cage studies of the caribfly (Dodson 1982, Hendrichs unpubl.
data), medfly (Prokopy and Hendrichs 1979), and the Queensland fruit fly,
Dacus tryoni (Froggatt) (Tychsen 1977), calling males were clearly ag-
gregated. In my study, males were observed calling both in aggregations
and singly. As many as 7-8 males were observed displaying and interacting
in an area no more than 0.25 m2. Perdomo et al. (1976) showed that male
caribfly sex pheromone was equally attractive to males and virgin females.
Fitt (1981) has remarked that the traditional male attractants of tephritids
such as Dacus opiliae Drew & Hardy, D. aquilonis (May), and D. tenuifascia
(May) also attract virgin females at precisely the time of day that they
are ready to mate, therefore acting in the manner of sex pheromones. If
calling males are normally found in aggregations, these results are less
anomalous than some authors have thought (Fitt 1981). If males enjoy
greater mating success in aggregations than when isolated, one would expect
the evolution of responses by males to cues that indicate the presence of
other displaying males. The chief of these would, of course, be sex pheromone.
These male-attractant effects of pheromones, which arise from the lek mating
system (lek = male sexual display aggregation), may have important impli-
cations for trapping and control of polyphagous tephritids.
341
Florida Entomologist 66(3)
Selection of males that tend to aggregate may occur if more females are
attracted per male to aggregations than to isolates or if females prefer to
mate after sampling a number of males (females sampling in male aggrega-
tions would reduce their energy cost and risk of predation). There is cur-
rently not enough information available to evaluate these 2 possibilities.
Working in a field cage where densities were higher and male aggregations
more prominent, Hendrichs (unpubl. data) found that an overwhelming
majority of matings occurred within rather than outside leks. He observed
virgin females in the vicinity of male aggregations for hours and sometimes
days prior to mating, suggesting female evaluation of males.
If, as seems likely, female caribflies usually mate with males that are
displaying and in a lek, the importance of male aggressive interactions to
reproductive success becomes apparent. To mate, a male must be in possession
of a leaf territory where he can puff and call, and must be in proximity to
other such males. He is subject to repeated intrusions by other males that he
must successfully repel in order to resume puffing and calling to attract
females. Thus, even though caribfly males are not fighting directly over a
resource that will in itself attract females (as do R. pomonella over apples
or R. indifferens Curran over cherries (Biggs 1972, AliNiazee 1974)), males
do compete for calling stations within leks which are prerequisites for the
attraction of females. Females, by observing leks or by responding prefer-
entially to territory-holding, puffing and calling males may on average mate
with more vigorous, dominant males. These traits should contribute to the
reproductive success of the females' offspring (Trivers 1972).
Finally, the significance of these findings for sterile-male release pro-
grams for the control of pest tephritids needs to be considered. A female
choosing a mate is acting in essentially the same way as an entomologist
concerned with the quality control of the sterile flies he is rearing and re-
leasing. Knowledge of the qualities selected by male competition and female
choice should allow the release of sterile males with a better chance of
competing with wild males. This study suggests that male aggressive ability
and factors such as size that contribute to it are likely to be important in
reproductive success of male caribflies. Quality control tests to assess male
aggressive ability may be needed.
Mention of a commercial or proprietary product does not constitute an
endorsement by the USDA.
ACKNOWLEDGMENTS
I would like to thank Andrew Duncan for allowing us to work at the
University of Florida, Agricultural Research and Education Center (AREC)
in Homestead, and Sandoz Farms for allowing us to wander among their
guava trees. R. M. Baranowski and R. W.,Swanson provided invaluable help
at the AREC. D. L. Chambers, J. C. Webb, and C. O. Calkins of this Labora-
tory and T. J. Walker of the University of Florida provided help, super-
vision, and criticism of the manuscript. Superior technical assistance was
provided by the late J. C. Benner, F. L. Lee and S. Masuda. J. Reiskind
identified Lyssomanes viridis. Elaine S. Turner typed the manuscript. Special
thanks are due to Jorge Hendrichs for many useful suggestions, discussions,
and acts of assistance.
342
September, 1983
Burk: Mating in Caribbean Fruit Flies
REFERENCES CITED
ALTMAN, J. 1974. Observational study of behavior: Sampling methods.
Behaviour 69: 227-67.
ALINIAZEE, M. T. 1974. The Western cherry fruit fly, Rhagoletis indifferens
(Diptera: Tephritidae). 2. Aggressive behavior. Canadian Ent. 106:
1201-4.
BIGGS, J. D. 1972. Aggressive behavior in the adult apple maggot (Diptera:
Tephritidae). Canadian Ent. 104: 349-53.
BURK, T. 1981. Signaling and sex in acalyptrate flies. Florida Ent. 64:
30-43.
CADE, W. 1980. Alternative male reproductive behaviors. Florida Ent. 63:
30-45.
DAWKINS, R. 1980. Good strategy or evolutionarily stable strategy? Pages
331-67 in G. W. Barlow and J. Silverberg (eds.), Sociobiology: Beyond
Nature/Nurture. Westview, Boulder, CO.
DODSON, G. N. 1978. Behavioral, anatomical, and physiological aspects of
reproduction in the Caribbean fruit fly, Anastrepha suspense (Loew).
M.Sc. thesis, University of Florida, Gainesville. 68 p.
1982. Mating and territoriality in wild Anastrepha suspense
(Diptera: Tephritidae) in field cages. J. Georgia Ent. Soc. 17: 189-200.
FITT, G. P. 1981. Responses by female Dacinae to "male" lures and their
relationship to patterns of mating behaviour and pheromone response.
Ent. exp. & appl. 29: 87-97.
MAIER, C. T., AND G. P. WALDBAUER. 1979. Dual mate-seeking strategies in
male syrphid flies (Diptera: Syrphidae). Ann. Ent. Soc. America 72:
54-61.
MYERS, H. S., B. D. BARRY, J. A. BURNSIDE, AND R. H. RHODE. 1976. Sperm
precedence in female apple maggots alternately mated to normal and
irradiated males. Ann. Ent. Soc. America 69: 39-41.
NATION, J. L. 1972. Courtship behavior and evidence for a sex attractant in
the male Caribbean fruit fly, Anastrepha suspense. Ann. Ent. Soc.
America 65: 1364-7.
PERDOMO, A. J. 1974. Sex and aggregation pheromone bioassays and mating
observations of the Caribbean fruit fly Anastrepha suspense (Loew)
under field conditions. Ph.D. dissertation, University of Florida,
Gainesville. 127 p.
- J. L. NATION, AND R. M. BARANOWSKI. 1976. Attraction of female
and male Caribbean fruit flies to food-baited and male-baited traps
under field conditions. Environ. Ent. 5: 1208-10.
PROKOPY, R. J. 1980. Mating behavior of frugivorous Tephritidae in nature.
Pages 37-46 in Proc. Symp. Fruit Fly Problems, XVI Int. Congr. Ent.,
Kyoto.
-- P. D. GREANY, AND D. L. CHAMBERS. 1977. Oviposition-deterring
pheromone in Anastrepha suspense. Environ. Ent. 6: 463-5.
- AND J. HENDRICHS. 1979. Mating behavior of Ceratitis capitata on a
field caged host tree. Ann. Ent. Soc. America 72: 642-8.
SMITH, D. C., AND R. J. PROKOPY. 1980. Mating behavior of Rhagoletis
pomonella (Diptera: Tephritidae) .VI. Site of early-season encounters.
Canadian Ent. 112: 585-90.
SUZUKI, Y., AND J. KOYAMA. 1981. Courtship behavior of the melon fly,
Dacus cucurbitae Coquillett (Diptera: Tephritidae). Appl. Ent. Zool.
16: 164-6.
SWANSON, R. W., AND R. M. BARANOWSKI. 1972. Host range and infestation
by the Caribbean fruit fly, Anastrepha suspense (Diptera: Tephri-
tidae) in South Florida. Proc. Florida St. Hort. Soc. 85: 271-4.
TRIVERS, R. L. 1972. Parental investment and sexual selection. Paves 136-17Q
343
Florida Entomologist 66 (3)
in Campbell, B. (ed.), Sexual Selection and the Descent of Man, 1871-
1971. Aldine, Chicago.
TYCHSEN, P. H. 1977. Mating behavior of the Queensland fruit fly, Dacus
tryoni, in field cages. J. Australian Ent. Soc. 16: 459-65.
WEBB, J. C., J. L. SHARP, D. L. CHAMBERS, J. J. McDow, AND J. C. BENNER.
1976. The analysis and identification of sounds produced by the male
Caribbean fruit fly, Anastrepha suspense (Loew). Ann. Ent. Soc.
America 69: 415-20.
-L -- a --
ECOLOGICAL NOTES ON LOWER RiO GRANDE VALLEY
AUGOCHLOROPSIS AND AGAPOSTEMON
(HYMENOPTERA: HALICTIDAE)
CHARLES C. PORTER1
ABSTRACT
Records are given from Hidalgo County, Texas for the Middle American
Neotropic Agapostemon nasutus Smith and the primarily North American
but Neotropic-derived Agapostemon melliventris Cresson, Agapostemon
texanus Cresson, and Augochloropsis metallica metallica (Fabricius). Re-
peated field studies during 2 years show that local populations of these
halictids differ among themselves in seasonal abundance, activity tempera-
tures, habitat preference, and in the flowers they visit for pollen and/or
nectar.
RESUME
Son citados del Condado de Hidalgo en Tejas los halictidos Agapostemon
nasutus Smith, especie neotropical-mesoamericana, y ademis Agapostemon
melliventris Cresson, A. texanus Cresson, Augochloropsis m. metallica
(Fabricus), species principalmente norteamericanas pero con afinidades
neotropicales.
Observaciones de campo repitidas atraves de 2 afios, demuestran que las
poblaciones locales de estas abejas difieren entire si en abundancia estacional,
relaciones termicas, ambientes naturales preferidos, y flores visitadas para
recoger o polen o nectar o ambas cosas.
Augochloropsis Cockerell and Agapostemon GuBrin include bright green
solitary bees that abound in much of the New World. Although not close
taxonomically (Eickwort 1969: 326), these genera often use the same habi-
tats and seem both to have originated in the Neotropics. Much taxonomic and
biological data have been compiled for the 2 genera by Eickwort (1969),
Mitchell (1960), and Roberts (1972). However, Lower Rio Grande Valley
populations of Augochloropsis and Agapostemon have remained largely un-
studied.
My work in south Texas was begun in 1973. Since December 1979, I have
been keeping records of hourly occurrence, activity temperatures, habitat
'Research Associate, Florida State Collection of Arthropods: Biology Department, Fordham
University, Bronx, NY 10458 USA.
September, 1983
344
Porter: Notes on Texas Halictids 345
selection, and plant relationships for most Hymenoptera collected in the
Lower Rio Grande Valley. Periods available for these studies have included
20 December 1979 to 20 January 1980, most of July and August 1980, 23 to 30
November 1980, 20 December 1980 to 16 January 1981, 10 to 17 March 1981,
4 to 18 June 1981, most of July and August 1981, 22 to 30 November 1981,
and 17 December 1981 to 6 January 1982. In the following contribution, I
present the results of this fieldwork as they apply to Augochloropsis and
Agapostemon.
MATERIAL STUDIED
Genus Augochloropsis Cockerell
1. Augochloropsis metallica metallica (Fabricius)
SPECIMENS EXAMINED. 47 9, 15 & : TEXAS, Hidalgo County, Bentsen Rio
Grande Valley State Park, 19, 1 S, 14-III-1981, 19, 16-III-1981, 2 9,
17-III-1981, 3 9, 18-III-1981, 1 $, 20-III-1981, 2 9, 21-III-1981, 1 9, 8-VI-
1981, 1 3, 30-VII-1980, 1 9, 1 3, 1-VIII-1981, 1 9, 7-VIII-1981, 1 3, 12-VIII-
1981, 1 9, 24-VIII-1981, 6 9, 25-VIII-1981, 1 9, 26-VIII-1981, 1 9, 24-XI-
1981; McAllen Botanical Gardens at McAllen, 2 9, 1 S, 1-1-1981, 3 9, 1 3,
3-1-1981, 1 9, 4-1-1981, 1 $, 5-1-1981, 2 8, 6-1-1981, 2 9, 15-III-1981, 1 3,
20-VII-1980, 1 $, 26-VII-1980, 2 9, 11-VIII-1980, 1 9, 12-VIII-1980, 1 9,
17-VIII-1980, 2 9, 23-VIII-1980, 2 9, 30-VIII-1981, 1 3, 26-XI-1981, 2 9,
30-XI-1980, 3 9, 26-XII-1980, 1 S, 27-XII-1980, 1 9, 28-XII-1979, 2 9, 28-
XII-1980, 2 9, 28-XII-1981.
GEOGRAPHIC DISTRIBUTION. This species ranges over the eastern and
southeastern United States west to Arizona and extends south in Middle
America to Panama.
FLOWER RECORDS. I have collected 55 A. metallica at flowers of 12
herbaceous and 3 woody plant species. These are listed below, according to
the number of specimens netted from each plant and to the months in which
samples were taken.
1. Condalia obovata Hook. (Rhamnaceae). 12 9,3 8, VII, VIII.
2. Heterotheca latifolia Buckl. (Compositae). 10 9,5 3, XI, XII, I.
3. Teucrium cubense Jacq. (Labiatae). 7 9,1 3, III.
4. Bumelia celastrina HBK. (Sapotaceae). 2 9,1 S, XI, XII.
5. Rivina humilis L. (Phytolaccaceae). 3 9, VII, VIII.
6. Solanum triquetrum Cav. (Solenaceae). 2 9, VIII.
7. Aster spinosus Benth. (Compositae). 1 9, XI.
8. Beloperone sp. (Acanthaceae). 1 9, I.
9. Croton sp. (Euphorbiaceae). 1 9, III.
10. Helianthus annuus L. (Compositae). 1 9, XII.
11. Ipomoea sp. (Convolvulaceae). 1 9, VIII.
12. Lppia alba Michx. (Verbenaceae), 1 9, VIII.
13. Mikania scandens L. (Compositae). 1 9, I.
14. Oenothera speciosa Nutt. (Onagraceae). 1 9, IT .
15. Stachys drummondii Benth. (Labiatae). 1 3, III.
HABITAT. This bee occurs in diverse natural and disturbed communities,
such as Celtis and Pithecellobium woods, thorn scrub, abandoned citrus
groves, and open fields. I have 33 records for woodlands and 22 for fields. Of
the 2 plants most often visited by A. metallica, Condalia obovata is an
Florida Entomologist 66(3)
arborescent component of the thorn scrub, while Heterotheca latifolia grows
in pioneering fields.
MONTHLY PHAENOLOGY. 6 9 and 5 $ in I, 11 9 and 2 $ in III, 1 9 in VI,
3 & in VII, 18 9 and 3 6 in VIII, 3 9 and 1 S in XI, and 8 9 and 1 $ in
XII.
These records suggest that the species becomes most abundant during late
summer and that it shows another major upsurge in early spring. It seems
rare through early and mid summer but is common in fall and early winter.
TEMPERATURE AND DIEL PERIODICITY. I have 48 temperature records for
A. metallica. It appears on sunny days at air temperatures between 21.5 and
37.00C. The median activity temperature is 27.50C.
Like other local Hymenoptera that fly throughout the year (e.g.,
Xylocopa, Porter 1981a or Campsomeris, Porter 1981b), this bee shows a
pronounced seasonal shift in diel periodicity (Table 1). I have 37 hourly
records for December, January, and March. All were registered between
1100 and 1600 CST, with 2 in the 1st hour, 9 in the 2nd, 5 in the 3rd, 9 in the
4th, and 12 in the 5th. During June to August, 25 hourly records were taken.
These extended from 0900 to 1400 CST, with 2 in the 1st hour, 5 in the 2nd,
10 in the 3rd, 4 in the 4th, and 4 in the 5th.
Genus Agapostemon GuBrin
1. Agapostemon nasutus Smith
SPECIMENS EXAMINED. 1 9, 1 $ : TEXAS, Hidalgo County, Bentsen Rio
Grande Valley State Park, 1 9, 14-VIII-1980, 1 $, 27-XII-1977.
GEOGRAPHIC DISTRIBUTION. A. nasutus ranges from south Texas to
Colombia and Peri.
FLOWER RECORDS AND HABITAT. My single female was netted from catkins
of Salix interior Rowlee at the edge of gallery woods along the Rio Grande
River.
MONTHLY PHAENOLOGY. The 2 south Texas records suggest a year-round
activity cycle. This is confirmed by Roberts (1972: 509), who states that A.
nasutus "has been collected in M6xico north of the Isthmus of Tehuantepec
in every month but January".
2. Agapostemon melliventris Cresson
SPECIMENS EXAMINED. 19 9, 95 8 : TEXAS, Hidalgo County, Bentsen Rio
Grande Valley State Park, 1 9, 10-1-1976, 1 $, 11-1-1976, 1 9, 1 8, 14-I-
1981, 1 9, 3 $, 15-1-1981, 1 9, 23-1-1977, 1 6, 5-VI-1973, 1 3, 10-VI-1977,
2 $, 14-VII-1981, 2 8, 5-VII-1981, 4 8, 17-VII-1981, 1 8, 18-VII-1980, 1 $,
22-VII-1980, 2 8, 24-VII-1980, 1 3, 25-VII-1980, 2 $, 31-VII-1981, 6 $, 1-
VIII-1981, 2 3, 3-VIII-1981, 6 8, 4-VIII-1981, 4 8, 5-VIII-1981, 5 8, 6-VIII-
1981, 1 8, 7-VIII-1981, 2 $, 10-VIII-1981, 1 9, 11-VIII-1981, 2 8, 12-VIII-
1981, 1 $, 13-VIII-1981, 4 8, 15-VIII-1981, 1 $, 15-VIII-1981, 2 8, 19-VIII-
1981, 1 $, 23-VIII-1977, 1 $, 23-VIII-1981, 1 $, 24-VIII-1979, 1 6, 26-VIII-
1981, 1 $, 28-VIII-1981, 1 8, 30-VIII-1977, 3 6, 29-XI-1980, 1 9,1 6, 27-
XII-1977, 2 8, 31-XIII-1980; McAllen Botanical Gardens at McAllen, 1 $,
11-1-1976, 1 3, 23-1-1977, 1 8, 16-30-V-1974, 1 8, 6-VI-1973, 2 8, 3-VII-
1980, 1 9, 12-VII-1981, 1 9, 3 11-VIII-1980, 2 8, 10-VIII-1980, 1 9, 1 ,
27-VIII-1973, 1 9, 30-VIII-1975, 2 9, 3-IX-1975, 2 8, 4-IX-1977, 1 9, 6-IX-
1975, 3 9, 7-IX-1975, 1 9, 8-IX-1975, 2 6, 27-XI-1980, 1 9, 20-XII-1981,
346
September, 1983
Porter: Notes on Texas Halictids
1 $, 24-XII-1977, 1 27-XII-1981, 1 9, 28-XII-1980, 1 9, 2 3, 28-XII-1981,
3 $, 29-XII-1977, 1 31-XII-1977.
GEOGRAPHIC DISTRIBUTION. A. melliventris extends over most of the
western United States and south into M6xico at least to the Isthmus of
Tehuantepec.
FLOWER RECORDS. I have collected 43 A. melliventris at flowers of 7
herbaceous and 3 woody plant species. These are listed below, according to
the number of specimens obtained and to the months in which samples were
taken.
1. Salix interior Rowlee (Salicaceae). 12 8, VII, VIII.
2. Condalia obovata Hook. (Rhamnaceae). 2 9,9 $, VIII, XII.
3. Rivina humilis L. (Phytolaccaceae). 4 9, VIII.
4. Mikania scandens L. (Compositae). 19, 3 8, XII, I.
5. Aster spinosus Benth. (Compositae). 3 3, XI.
6. Heterotheca latifolia Buckl. (Compositae). 1 9,2 8, XI, XII.
7. Teucrium cubense Jacq. (Labiatae). 3 9, XII.
8. Bumelia celastrina HBK. (Sapotaceae). 1 9, XI.
9. Helianthus annuus L. (Compositae). 1 $, I.
10. Palafoxia texana DC. (Compositae). 1 9, VII.
HABITAT. Agapostemon melliventris frequents many local habitats, such
as pioneering fields, thorn scrub, and semihumid forest. Both sexes visit
flowers mostly in exposed, sunny places. During late July and August, males
suddenly become abundant. The collection records given above reflect only a
small percent of this dramatic increase. They fly by the hundreds near the
ground amid tall grass, in herbaceous undergrowth, beneath dense bushes
and in other shaded or semi-shaded sites. This species consequently often
feeds and forages in the open but resorts to woodland for other (presumably
reproductive) functions.
MONTHLY PHAENOLOGY. 4 9 and 7 & in I, 1 & in V, 3 & in VI, 1 9 and
17 8 in VII, 4 9 and 48 $ in VIII, 7 9 and 2 $ in IX, 5 $ in XI, and 4 9
and 11 S in XII.
Local populations of this halictid, as far as studied, emerge during May,
become very abundant by August and September, and remain fairly common
through December and January. I have not taken A. melliventris in March
(at least 1 week's collecting during every year since 1974). Females have
been found only during July, August, September, December, and January.
The phaenologic trend documented by sampling between July and the 1st
week of September suggests that female activity may peak from mid Septem-
ber to October.
TEMPERATURE AND DIEL PERIODICITY. This bee flies on sunny days. I have
38 temperature records, which show that its activity range lies between 19
and 36'C. The median activity temperature is 290C.
I have 84 hourly records for A. melliventris, registered in July, August,
November, December, and January. During the summer months, it appeared
between 0900 and 1400 CST, with 7 specimens in the 1st hour, 13 in the 2nd,
21 in the 3rd, 11 in the 4th, and 13 in the 5th. Late fall and early winter
collecting revealed a 1200 to 1600 CST flight period, with 3 records for the
1st hour, 7 for the 2nd, 7 for the 3rd, and 2 in the 4th.
3. Agapostemon texanus Cresson
SPECIMENS EXAMINED. 34 9, 71 : TEXAS, Hidalgo County, Bentsen Rio
347
Florida Entomologist 66 (3)
Grande Valley State Park, 1 3, 2-1-1981, 1 8, 5-1-1978, 2 9, 6-1-1978, 4 $,
14-1-1981, 3 $, 15-1-1981, 1 $, 20-V-1977, 1 3, 7-VI-1973, 1 9, 12-VI-1981,
1 3, 13-VI-1979, 1 9, 9-VII-1981, 1 9, 24-VII-1980, 1 3, 28-VII-1980, 1 3,
5-VIII-1981, 2 3, 10-VIII-1981, 3 8, 12-VIII-1981, 1 3, 15-VIII-1981, 2 8,
24-VIII-1979, 2 8, 28-XI-1980, 2 9, 1 8, 29-XI-1980, 1 8, 23-XII-1975, 2 3,
27-XII-1977, 1 9,3 $, 3-XII-1980; McAllen Botanical Gardens at McAllen,
1 3, 1-1-1981, 6 9, 2 S, 2-1-1982, 4 $, 3-1-1981, 2 $, 4-1-1981, 2 $, 6-1-1981,
1 9, 12-1-1974, 1 9, 18-1-1977, 1 9, 11-III-1979, 3 9, 17-24-III-1974, 1 9,
21-III-1976, 1 9, 1-13-VI-1973, 1 9, 7-VI-1979, 1 $, 14-VI-1981, 2 8, 3-VII-
1980, 1 8, 4-VII-1980, 1 9, 2 8, 6-VII-1980, 1 9, 12-VII-1981, 1 $, 19-VII-
1981, 2 8, 27-VIII-1977, 1 S, 28-VIII-1977, 1 3, 31-VIII-1976, 1 9, 6 3,
1-8-IX-1973, 3 3, 27-XI-1980, 1 9, 1 8, 20-XII-1973, 1 8, 23-XII-1975, 4 9,
1 8, 26-XII-1980, 2 9, 27-XII-1980, 3 8, 29-XII-1975, 2 8, 30-XII-1975,
1 9, 31-XII-1975, 1 9, 31-XII-1981.
GEOGRAPHIC DISTRIBUTION. This species ranges over most of temperate,
subtropical, and tropical North and Middle America.
FLOWER RECORDS. I have collected 78 A. texanus at flowers of 9 herbaceous
and 1 woody plant species. These are listed below, according to the number
of specimens obtained from each plant and to the months in which samples
were taken.
1. Helianthus annuus L. (Compositae). 16 9, 34 $, VI, VII, XII, I.
2. Heterotheca latifolia Buckl. (Compositae). 2 9, 6 $, XI, XII, I.
3. Aster spinosus Benth. (Compositae). 1 9, 3 3, XI.
4. Mikania scandens L. (Compositae). 1 9, 3 $, XII, I.
5. Verbesina encelioides Cav. (Compositae). 3 9, 1 $, XI, I.
6. Pluchea purpurascens Sw. (Compositae). 2 9, 1 6, VII.
7. Teucrium cubense Jacq. (Labiatae). 2 S, I.
8. Lippia alba Mill. (Verbenaceae). 1 8, XII.
9. Palafoxia texana DC. (Compositae). 1 9, VII.
10. Sonchus sp. (Compositae). 1 8, I.
HABITAT. Local populations of this bee frequent principally fields and
open places in scrub or woodland. I have 69 habitat records, of which 7 are
for open woods, 4 from extensive woodland clearings, and 58 from fields and
abandoned citrus groves. Occasionally, I have swept males of A. texanus
from shaded undergrowth where males of A. melliventris were swarming.
However, during the times available for fieldwork, I have never observed
large aggregations of A. texanus males in any habitat.
MONTHLY PHAENOLOGY. 10 9 and 20 6 in I, 5 6 in III, 1 $ in V, 3 9 and
3 6 in VI, 4 9 and 7 S in VII, 14 $ in VIII, 1 9 and 6 6 in IX, 2 9 and
6 8 in XI, and 9 9 and 14 8 in XII.
A. texanus probably occurs throughout the year in south Texas. It peaks
in December and January, becomes scarce by March and May, and then
increases markedly during summer and fall. I have found no females in
March, May, and August. Of the 34 fernales collected, 19 were obtained in
December and January.
Roberts (1972: 534) reported data on seasonal occurrence of populations
from other parts of this species' extensive range. In British Columbia it has
been collected from May to October, in Kansas from April through Novem-
ber, and in M6xico throughout the year. Roberts also noted geographic varia-
tion in sexual phaenology, with males occurring in British Columbia only
September, 1983
348
Porter: Notes on Texas Halictids
from "July to October", in Kansas from "April (very rare) and from June
to October", and in M6xico only from "April and June through September".
TEMPERATURE AND DIEL PERIODICITY. Agapostemon texanus is active
under clear or partly cloudy conditions. My 72 temperature records for this
bee range from 16 to 370C. The median activity temperature is 240C.
Hourly occurrence was noted for 78 specimens collected during June,
July, August, November, December, and January. In summer, it appeared
between 0700 and 1500 CST, with 1 record for the 1st hour, 2 for the 2nd, 2
for the 3rd, 1 for the 4th, 5 for the 5th, 7 for the 6th, 1 for the 8th, and 1
for the 9th. From November to January, this bee was active between 1100
and 1600, with 5 records noted in the 1st hour, 10 in the 2nd, 21 in the 3rd, 9
in the 4th, and 12 in the 5th.
CONCLUSIONS
Augochloropsis metallica is a North and Middle American species that
belongs to a genus which is best developed in semihumid to humid parts of
the Neotropics. The Agapostemon generic group also seems to be a Neotropic
evolutionary product but shows comparatively xeric habitat proclivities.
Agapostemon melliventris and A. texanus are North and Middle American
species of apparent Sonoran origin. A. melliventris remains centered in the
Sonoran Biogeographic Province (southwestern United States and northern
M6xico). A. texanus, perhaps because of its notable cold tolerance, has spread
far beyond the Sonoran region to occupy most of the United States and
southern Canada. In contrast, A. nasutus ranges from the tip of Texas to the
Peruvian Coastal Desert and belongs to a Middle American Neotropic radia-
tion. Except for A. nasutus, all of these species are extremely common in the
Lower Rio Grande Valley.
Both Augochloropsis and Agapostemon are polylectic bees. However, the
3 common local species diverge considerably in flower preference (Table 2).
Augochloropsis metallica visits 15 species of plants. Condalia obovata,
Heterotheca latifolia, and Teucrium cubense account for 38 of the 55 speci-
mens collected on flowers. Agapostemon melliventris furnishes 43 specimens
taken on 10 plant species. Salix interior (12 8 ) and Condalia obovata (2 9,
8 $ ) provide the greatest number of floral records. These plants seem to be
visited for nectar only. Scattered females of A. melliventris appear on Rivina
humilis, Teucrium cubense, Heterotheca latifolia, Bumelia celastrina, and
Palafoxia texana. To determine its major pollen sources, this bee should be
studied during September and October, the months of predicted maximum
female activity. Agapostemon texanus visits 10 local plant species, of which
8 are Compositae. Its main pollen and nectar source is Helianthus annuus
(16 9 and 34 8). I have no more than 2 9 and 6 $ from any other species
of flower and only 3 specimens from non-composites (2 $ on Teucrium
cubense and 1 $ on Lippia alba).
Partly according to floral selection, the halictids studied differ in several
habitat factors. Almost 70% of the Augochloropsis metallica are from
natural thorn scrub and woods, while 30% are from open fields or orchards.
This bee ranges from low herbs (Teucrium cubense) to plants in the upper
strata of its communities (Condalia obovata). Agapostemon melliventris
predor inmates in woods, but commonly visits flowers in open fields. Like
Augochloropsis, it occurs regularly in all strata of its habitats. When seeking
349
Florida Entomologist 66 (3)
September, 1983
RECORDS OF HOURLY ACTIVITY (CST) FOR Augochloropsis metallica
(MT), Agapostemon melliventris (ML) AND Agapostemon
texanus (TX).
Jan. March June July Aug. Nov. Dec.
TX-1
MT-3
TX-2
MT-1
ML-5
TX-6
ML-1
TX-5
MT-7
TX-3
MT-2
MT-3
MT-2
MT-5
MT-1
MT-1
TX-1
TX-1
TX-1
TX-2
MT-1
ML-4
TX-2
ML-2
TX-1
MT-1
ML-5
TX-3
MT-1
ML-3
TX-1
TX-1
0700-0800
0800-0900
0900-1000
1000-1100
1100-1200
1200-1300
1300-1400
1400-1500
1500-1600
MT-1
ML-3
MT-5
ML-11
MT-8
ML-16
TX-1
MT-3
ML-11
TX-6
MT-4
ML-10
TX-1
TX-1
MT-2
TX-2
ML-2
TX-1
-
TX-3
MT-1
ML-3
TX-6
MT-2
TX-14
- MT-4
ML-4 ML-2
- TX-4
MT-2
TX-3
MT-2
ML-2
TX-6
nectar, A. melliventris
often ascends to overstory
plants (51% of all speci-
mens on Condalia obovata and Salix interior). Agapostemon texanus differs
in spatial proclivities from the 2 proceeding species. Only 14% of my records
are from woods, whereas 86% correspond to fields and orchards. A. texanus
inhabits almost exclusively herbaceous strata, visiting plants of 2 m or less
in height.
These common south Texas Halictidae also manifest divergent monthly
phaenologies (Table 3). Augochloropsis metallica is common in winter, re-
mains abundant in March, and shows another peak in August. Agapostemon
melliventris gives a similar phaenologic pattern, except that it is absent in
March and has a more pronounced summer peak. In contrast, Agapostemon
texanus has its maximum abundance during December and January, shows
a secondary upsurge in July and August, but is comparatively scarce during
other months sampled.
Local Augochloropsis and Agapostemon also have different threshold and
median activity temperatures. Augochloropsis metallica shows a threshold of
21.50C and a median of 27.50C. The threshold for Agapostemon melliventris
is 190C and the median 290C. Agapostemon texanus shows the greatest
350
TABLE 1.
TX-1
Porter: Notes on Texas Halictids
TABLE 2. FLOWER RECORDS FOR LOWER
AND Agapostemon.
RIo GRAND VALLEY Augochloropsis
A. A. A. A.
metallica nasutus melliventris texanus
Aster spinosus 1 3 4
Beloperone sp. 1 --
Bumelia celastrina 3 1 -
Condalia obovata 15 10
Croton sp. 1 --
Helianthus annuus 1 1 50
Heterotheca latifolia 15 3 7
Ipomoea sp. 1 -
Lippia alba 1 1
Mikania scandens 1 4 4
Oenothera speciosa 1 -
Palafoxia texana 1 1
Pluchea purpurascens 3
Rivina humilis 3 4
Salix interior 1 12
Solanum triquetrum 1 -
Sonchus sp. 1
Stachys drummondii 1 --
Teucrium cubense 8 4 2
Verbesina encelioides 3
Total species of plants
visited/bee species 15 1 10 10
climatic tolerance. Even on partly cloudy days, it begins to fly at 160C, and
its median activity temperature is 24C. All 3 species remain active up to at
least 370C and seem to appear in nearly equal numbers at any temperature
above threshold. These thermic data seem reflected in diel periodicity (Table
1). During summer, all species fly primarily in the morning, but in winter
shift most activity to the afternoon. Augochloropsis metallica and
Agapostemon melliventris show similar daily flight patterns, but Aga-
postemon texanus begins summer activity 2 hours earlier than the other
species and emerges on winter days approximately 1 hour earlier.
The ecological observations made above apply only to south Texas popula-
tions of these wide-ranging Halictidae. Agapostemon, in particular, shows
much ecological variation along geographic lines. Roberts (1972: 534) re-
marked, for example, that "A. texanus has been found in 70 of the 116 major
United States plant communities" and that "considering the lack of ap-
parent climatic, biotic, or edaphic factors correlated with its distribution",
the distribution appears "inexplicable". Long-term, geographically compre-
hensive biosystematic analysis of these 'wide-ranging, generally adapted
taxa will be required to ascertain the exact relationships of their spatially
distant, adaptively disparate, and taxonomically variable populations.
COLLECTIONS
This study is based on collections made by the author. The material
covered has been divided between the Florida State Collection of Arthropods
352
September, 1983
Florida Entomologist 66 (3)
cq cq1 MI
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a
P4
Sal 00
LO 00 crt--
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C-
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Porter: Notes on Texas Halictids 353
(Division of Plant Industry, P.O. Box 1269, Gainesville, FL 32602) and the
author's private collection (301 N. 39th St., McAllen, TX 78501).
ACKNOWLEDGMENTS
Fieldwork in Texas was supported during 1980 by a Faculty Fellowship
from Fordham University, during 1973-78 and in 1979 and 1981 by grants
from the Committee for Research and Exploration of the National Geo-
graphic Society, and from 1976-77 by United States National Science
Foundation Grant DEB-75-22426. Collecting in Bentsen Rio Grande Valley
State Park was done under permits issued by the Texas Parks and Wildlife
Department (current permit number 1-82). Mr. Anthony F. Cerbone of
Texas A & M University assisted during some of the fieldwork contributing
to this research.
Contribution No. 563, Bureau of Entomology, Division of Plant Industry.
Florida Department of Agriculture and Consumer Services, Gainesville, FL
32602 USA.
REFERENCES CITED
EICKWORT, G. C. 1969. A comparative morphological study and generic
revision of the augochlorine bees (Hymenoptera: Halictidae). Univ.
Kansas Sci. Bull. 48: 325-524.
MITCHELL, T. B. 1960. Bees of the eastern United States. Volume I. North
Carolina Agric. Exper. Sta. Tech. Bull. 141: 1-538.
PORTER, C. C. 1981a. Ecological notes on Lower Rio Grande Valley Xylocopa
(Hymenoptera: Anthophoridae). Florida Ent. 64: 175-82.
S1981b. Scoliidae (Hymenoptera) of the Lower Rio Grande Valley.
Florida Ent. 64: 441-53.
ROBERTS, R. B. 1972. Revision of the bee genus Agapostemon (Hymenoptera:
Halictidae). Univ. Kansas Sci. Bull. 49: 437-590.
EIPHOSOMA DENTATOR (FABRICIUS) IN FLORIDA
(HYMENOPTERA: ICHNEUMONIDAE)
CHARLES C. PORTER1
ABSTRACT
Eiphosoma dentator (Fabricius) previously has been known from western
North America south to Argentina, and from Cuba and other Caribbean
islands. The first Florida specimens (118 9 ) were taken in 1979-80 by
Malaise traps on Fleming Key in Monroe County. This species differs from
other Eiphosoma by its pale-banded hind tibia, hyaline fore wing apex,
shining propodeum, complete areolet, and distally effaced cubital and sub-
discoidal veins. E. dentator prefers arid habitats and may have moved east
to Florida during Pleistocene xerothermic episodes. Its maximum adult
activity occurs in summer, while most Neotropic ichneumonids peak between
fall and spring.
'Research Associate, Florida State Collection of Arthropods; Biology Department, Fordham
University, Bronx, NY 10458 USA.
354 Florida Entomologist 66 (3) September, 1983
RESUME
Eiphosoma dentator (Fabricius) ha sido reportado previamente de
muchos lugares entire el oeste de Norteamerica y la Argentina, incluyendo
Cuba y otras islas del Mar Caribe. Los primeros ejemplares floridenses
(118 9) provinieron de Cayo Fleming en el Condado de Monroe y fueron
colectados durante 1979-80 en trampas de Malaise. La especie se distingue
de otros Eiphosoma por tener un anillo blancuzco en la tibia posterior, por
sus alas apicalmente transparentes, el propodeo reluciente, la segunda celda
cubital complete, y por sus venas cubital y subdiscoidal muy d4biles dis-
talmente. E. dentator prefiere ambientes Aridos y posiblemente lleg6 a Flor-
ida, desde el oeste, durante algdn episodio xerot6rmico del Pleistoceno.
Alcanza la mayor actividad adulta durante el verano, mientras que muchas
Ichneumonidae neotropicales prefieren los meses mAs frescos.
Through courtesy of Dr. Lionel A. Stange, Taxonomic Entomologist at
the Florida State Collection of Arthropods, I have received for study 118
Eiphosoma dentator (Fabricius). This material was collected by Malaise
traps during 1979-80 on Fleming Key in Monroe County, Florida. These are
the first records of E. dentator from the eastern United States. Previous
authors (Dasch 1979, Townes 1966) have reported the species only in
western North America (British Columbia to south Texas), in the West
Indies (Cuba, Grenada, Puerto Rico, St. Croix, St. Vincent, and Trinidad),
and in the mainland Neotropics from M6xico to Argentina.
Eiphosoma Cresson belongs to the Subfamily Cremastinae. Most
Eiphosoma are medium-sized to large, delicately slender wasps with an
unusually long and strongly compressed gaster. They often have the head
and mesosoma black and yellow and the gaster fulvous or ferruginous with
variably developed dusky areas. Eiphosoma differs from most other
ichneumonids in possessing a ventral preapical tooth on the hind femur. This
character also appears in the sympatric cremastines Pristomerus Curtis and
Xiphosomella Sz6pligeti. Eiphosoma may be separated from these genera be-
cause it lacks the thyridium (a pale, opaque, slightly depressed spot located
dorso-laterally on each side of the 2nd gastric tergite at some point between
the spiracle and base).
The genus contains 9 described North American species and at least 29
Neotropic species (Dasch 1979: 65). During my own Neotropic collecting,
I have taken it in lowland rainforest, tropical and subtropical deciduous
forest, torrid thorn scrub, desert scrub, and even in deciduous woods at
altitudes of up to 2000 m (Porter 1975b: 233). Eiphosoma tends to prefer
open woods, fields, and other sunny habitats that are avoided by many
ichneumonids. In the tropics and subtropics it continues active throughout
the summer. Most Neotropic Ichneumonidae peak in fall, spring, or winter
and decline in summer.
In this contribution, I offer notes on taxonomy, phaenology, zoogeography,
and hosts of E. dentator.
Eiphosoma dentator (Fabricius)
(Fig. 2, 3, 4)
MATERIAL EXAMINED. 118 9: FLORIDA, Monroe County, Fleming Key, 1,
10-1-1980; 1, 11-13-1-1980; 1, 15-1-1980; 1, 17-1-1980; 2, 22-1-1980; 1, 27-I-
Porter: Eiphosoma dentator in Florida
1980; 2, 9-II-1980; 1, 15-17-II-1980; 2, 22-23-II-1980; 1, 24-II-1980; 1, 27-II-
1980; 1, 8-9-III-1980; 1, 10-III-1980; 1, 13-III-1980; 1, 15-16-III-1980; 1, 17-
III-1980; 1, 22-23-III-1980; 2, 30-III-1980; 2, 31-III-1980; 1, 12-13-IV-1980;
1, 18-IV-1980; 1, 17-18-V-1980; 1, 20-V-1980; 1, 21-V-1979; 1, 22-V-1979; 1,
22-23-V-1980; 1, 24-V-1979; 1, 25-V-1979; 1, 29-V-1979; 1, 29-V-1980; 2, 30-
V-1979; 3, 4-VI-1979; 1, 6-VI-1980; 1, 6-VI-1979, 2, 8-10-VI-1979; 1, 11-VI-
1979; 1, 12-VI-1979; 1, 13-VI-1979 2, 14-VI-1979; 1, 18-VI-1979; 1, 19-VI-
1979; 1, 20-VI-1979; 1, 3-4-VII-1979; 2, 16-VII-1979; 3, 17-VII-1979; 2, 18-
VII-1979; 2, 19-VII-1979; 1, 20-VII-1979; 1, 23-VII-1979; 2, 24-VII-1979; 1,
26-VII-1979; 9, 30-VII-1979 4, 31-VII-1979; G, 2-3-VIII-1979; 6, 7-VIII-1979;
2, 9-VIII-1979; 3, 10-VIII-80; 6, 13-VIII-1979; 1, 14-VIII-1979; 2, 16-VIII-
1979; 17-VIII-1979; 1, 6-XI-1979; 1, 15-XI-1979; 1, 16-19-XI-1979; 1, 19-XI-
1979; 1, 20-XI-1979; 1, 5-XII-1979; 1, 6-XII-1979; 1, 8-9-XII-1979; 1, 10-XII-
1979; 2, 21-25-XII-1979.
All specimens listed above were collected in Malaise traps operated under
the supervision of Dr. Howard V. Weems, Jr., Curator of the Florida State
Arthropod Collection. It is remarkable that only females were obtained.
Dasch (1979: 79) found both sexes almost equally abundant (39 9, 36 8)
in the material he examined for his revision of the Nearctic Cremastinae.
TAXONOMY. The following combination of characters separates E. dentator
from all known Eiphosoma: mesosoma black with profuse yellow markings;
fore wing hyaline throughout (Fig. 2); hind tibia with a prominent dull
white median annulus (Fig. 4); clypeus 1.5-1.7 as wide at apex as high from
level of tentorial pits to middle of apical margin; malar space 0.53-0.83 as
long as basal width of mandible; hypostomal carina weakly flange-like;
mesoscutum and mesopleuron in great part with moderately large, sharp,
dense punctures that mostly are separated by a little more to a little less
than their diameters; propodeum shining, mostly with dense, sharp punc-
tures, the median depression punctate and wrinkled but not shagreened, the
median and lateral longitudinal carinae absent, and the pleural carina mostly
obsolete; sheathed portion of ovipositor 0.61-0.69 as long as fore wing; length
of fore wing 5-6 mm, areolet well developed, cubitus and subdiscoideus mostly
unsclerotized distad of 2nd recurrent vein; hind tarsus weakly compressed,
its claws with 4 minute teeth.
The only other Eiphosoma captured on Fleming Key was a single a of
E. nigrovittatum Cresson (Fig. 1), which entered a Malaise trap on 28-XII-
1979. This species differs from E. dentator in its unbanded hind tibia, apically
brownish fore wing, broader clypeus (2.3 as wide at apex as high from level
of tentorial pits to middle of apical margin), shagreened propodeal de-
pression, absent areolet, and fully sclerotized cubital and subdiscoidal veins.
Dasch (1979: 76) recorded E. nigrovittatum from Puerto Rico, Cuba, and
from south Florida (Key Largo in Monroe County and an unspecified locality
in Dade County).
MONTHLY PHAENOLOGY. 7 in 1-1980 (0.78 specimens/day), 7 in II-1980
(0.78 specimens/day), 10 in III-1980 (0.71 specimens/day), 2 in IV-1980
(0.50 specimens/day), 7 in V-1979 (0.86 specimens/day), 4 in V-1980 (0.44
specimens/day), 14 in VI-1979 (1.3 specimens/day), 1 in VI-1980 (1 speci-
men/day), 28 in VII-1979 (2.4 specimens/day), 27 in VIII-1979 (3.0 speci-
mens/day), no records for IX and X, 5 in XI-1979 (0.71 specimens/day), 6
in XII-1979 (0.71 specimens/day).
355
Florida Entomologist 66 (3)
September, 1983
356
1
4
Porter: Eiphosoma dentator in Florida
The preceding data show that Fleming Key populations of E. dentator
peak in July and August but apparently fly in reduced abundance during
most other months. Such year-round activity is not surprising, because the
Florida Keys enjoy an almost tropical climate. The Lower Keys have no
killing frost. At Key West, the July average temperature is 28.50C, the
January average is 210C, about 700 mm of rain accumulate from May to
October, and another 250 mm falls between November and April (Byers
1930: 225). The summer maximum shown on Fleming Key by E. dentator
coincides with phaenology of this and other Eiphosoma which I have observed
in hot, lowland Neotropic areas, such as the Lower Rio Grande Valley of
Texas, Santa Cruz in Bolivia, and northwest Argentine Chaco and Chaco
Forest.
In contrast, most common Fleming Key ichneumonids reach maximum
abundance between late fall and early spring. For example, traps operated
on this island from April to November yielded only 30 specimens of the
ephialtine, Calliephialtes ferrugineus Cushman, and but 11 of the ichneu-
monine, Paraditremops albipectus (Brull). Trapping between December
and March produced 75 C. ferrugineus and 66 P. albipectus. This same
invernally maximal and aestivally minimal seasonal pattern is displayed by
numerous ichneumonid faunas from diverse parts of the southern United
States and Latin America (Porter 1977: 81-6).
ZOOGEOGRAPHY. Eiphosoma dentator occupies a Neotropic and Sonoran
distribution in the southern United States and tropical America. It appears
to be the most xerophilous Eiphosoma and thrives in south Texas, the
Argentine Subandean Desert, and other dessicating environments (Porter
1975b: 233).
Spencer and Stegmaier (1973: 13) mentioned that "during a past period
of low rainfall a prairie type flora, such as is found today in Texas and
Arizona, was able to extend its range to the east, and remnants of this flora
reflecting dry conditions still exist on parts of the west coast" (of Florida)
"and the Lower Keys, such as Big Pine Key. Cactus thicket is the typical
plant association of these areas." Such "periods of low rainfall" occurred
both in xerothermic Pleistocene glacial minima and at times during the in-
creasingly arid, climatically unsettled late Tertiary. They allowed eastward
movement of southwestern biota and probably account for the contemporary
presence in Florida of relict species belonging to such Sonoran ichneumonid
genera as Compsocryptus Ashmead and Lanugo Townes (Porter 1975: 254).
Eiphosoma dentator thus could have reached Florida from the Sonoran
region via Texas and the Gulf states by the same mechanisms postulated
above for Compsocryptus and Lanugo. On the other hand, hurricanes may
have blown it across the Florida Straits from Cuba or it may have been
Fig. 1-4. 1) Eiphosoma nigrovittatum, $, Fleming Key, Florida. Fore
wing beyond basal vein, showing venation and infuscate apex. 2) Eiphosoma
dentator, 9, Fleming Key, Florida. Fore wing beyond basal vein, showing
venation and lack of apical infuscation. 3) Eiphosoma dentator, 9, Fleming
Key, Florida. Lateral view of hind femur, showing color pattern and the
preapical ventral tooth. 4) Eiphosoma dentator, 9, Fleming Key, Florida.
Lateral view of hind tibia showing pale annulus and other features of color
pattern.
357
Florida Entomologist 66 (3)
accidentally introduced into the Keys by man. More collecting on the Lower
Keys, which remain poorly sampled even for conspicuous insects like
Lepidoptera (Kimball 1965: 4), should help clarify the distributional and
ecological status of E. dentator.
HOSTS. This parasitoid has been reared from several injurious Lepi-
doptera, whose ranges include south Florida. It attacks Lineodes interga
(Zeller), a pyralid pest of eggplant and other Solanaceae. It also parasitizes
the phycitid, Ufa rubedinella (Zeller), an enemy of lima beans and black-
eyed peas. These host records were attested by Dasch (1979: 80).
COLLECTIONS
This study is based entirely on material deposited in the Florida State
Collection of Arthropods (Division of Plant Industry, P. O. Box 1269, Gaines-
ville, FL 32602.
ACKNOWLEDGMENTS
,Mr. Harold A. Denmark, Dr. Howard V. Weems, Jr., and Dr. Lionel A.
Stange of the Florida Department of Agriculture and Consumer Services at
Gainesville arranged for loan of ichneumonids collected on Fleming Key. I
am also deeply grateful to these friends and colleagues for their generous
support of my own work in Florida.
Contribution No. 532, Bureau of Entomology, Division of Plant Industry,
Florida Department of Agriculture and Consumer Services, Gainesville, FL
32602 USA.
REFERENCES CITED
BYERS, C. F. 1930. A contribution to the knowledge of Florida Odonata.
University of Florida Publ., Biological Science Series 1: 1-327.
DASCH, C. E. 1979. Ichneumon-flies of America north of M6xico: 8. Sub-
family Cremastinae. Mem. American Ent. Inst. 29: 1-702.
KIMBALL, C. P. 1965. Lepidoptera of Florida. Arthropods of Florida and
neighboring land areas 1: 1-363. Division of Plant Industry, Gaines-
ville.
PORTER, C. 1975a. A new Floridian Polycyrtidea with comments on zoo-
geography of Florida Mesostenini. Florida Ent. 58: 247-56.
1975b. Relaciones zoogeogrdficas y origen de la fauna de Ichneu-
monidae en la provincia biogeografica del monte del noroeste argentino.
Acta Zool. Lilloana 31: 175-250.
1977. Ecology, zoogeography, and taxonomy of the Lower Rio
Grande Valley mesostenines. Psyche 84: 28-91.
Spencer, K. A., and C. E. Stegmaier, Jr. 1973. Agromyzidae of Florida.
Arthropods of Florida and neighboring land areas 7: 1-205. Division
of Plant Industry, Gainesville.
TOWNES, H. K. 1966. A catalog and reclassification of the Neotropic
Ichneumonidae. Mem. American Ent. Inst. 8: 1-367.
358
September, 1983
Scientific Notes
SCIENTIFIC NOTES
PLUMERIA RUBRA: A NEW HOST PLANT RECORD OF THE FALL
ARMYWORM (LEPIDOPTERA: NOCTUIDAE) IN PUERTO RICO-
The fall armyworm, Spodoptera frugiperda (J. E. Smith), is a major agri-
cultural pest in Puerto Rico, the United States, and other countries. On the
Island it has been reported on weeds, pastures, and crops (Martorell. 1975.
Annotated Food Plant Catalog of the Insects of Puerto Rico. Agric. Exp.
Sta., Univ. Puerto Rico).
During July 1981, while examining leaves of the ubiquitous, exotic, orna-
mental tree Plumeria rubra L. (Apocynaceae), locally called "aleli" or "ramo
de novia", several egg clusters of S. frugiperda were found. Larvae of S.
frugiperda were fed in the laboratory with P. rubra leaf pieces until adults
emerged. This is the first record of the fall armyworm on Apocynaceae for
Puerto Rico although other lepidopterans, as well as other insects, have been
previously reported for P. rubra (Martorell, op. cit.).
Dr. R. W. Poole (Research Scientist, Insect Identification and Beneficial
Insect Introduction, ARS, USDA, Maryland) kindly identified the specimens.
Dr. S. Medina (Univ. P.R., Botanical Garden, Rio Piedras) and Dr. Jos6 A.
Mari Mutt (Univ. P.R., Mayaguez Campus, Biology Department) provided
comments to the manuscript.-J. A. SANTIAGO-BLAY, Univ. Puerto Rico,
Agricultural Experiment Station, Crop Protection Department, Venezuela
Contract Station, Rio Piedras, Puerto Rico (Present Address: Calle 33 ZG-9
Riverview, Bayam6n, Puerto Rico 00619 USA.)
OPPORTUNISTIC BLOOD-FEEDING ON EGG-LAYING SEA TURTLES
BY SALT MARSH MOSQUITOES (DIPTERA: CULICIDAE)-Few
mosquito species are parochial in their choice of vertebrate hosts for blood
meals. In fact, the opposite appears to be true. Most mosquitoes feed on a
range of host animals; many even feed readily on vertebrates from different
classes. Reasons for the diversity of host-feeding patterns observed among
mosquitoes in nature are not clear, and are reviewed elsewhere (Edman and
Kale 1971. Ann. Ent. Soc. America 64: 513-6; Tempelis 1975. J. Med. Ent.
11: 635-53). The plasticity of blood-feeding behavior by some mosquito
species is remarkable and unappreciated. The purpose of this note is to
report field observations of blood-feeding by the salt marsh mosquito Aedes
taeniorhynchus (Wiedemann) on egg-laying Atlantic loggerhead sea turtles,
Caretta caretta caretta (L.).
On 10 July 1982, between 2230-2330 h, 3 turtles were observed laying eggs
along the dune line of Floridana Beach, Brevard County, Florida. Weather
conditions that evening were: skies-clear, temperature-820F, RH-90%,
wind-calm. We observed many Ae. taeniorhynchus blood-feeding on the
front and back flippers, tail and neck region of all egg-laying turtles. We
estimated the biting count to be from 25 to 30 per minute on humans. Turtles
did not respond to mosquito feeding in any sort of defensive manner.
Mosquitoes engorged quickly and departed toward vegetation along the top
of the dune. Turtles took approximately 30 min to complete egg-laying.
In a comprehensive review of Aedes spp. blood-feeding in nature. Edman
359
Florida Entomologist 66(3)
(1971. J. Med. Ent. 8: 687-95) reported that Ae. taeniorhynchus fed pre-
dominantly on mammals, with less than 1% feeding on reptiles or am-
phibians. Engorged Ae. taeniorhynchus in Edman's study were collected
from 3 different habitats not including the coastal sand dune area. Had
Edman collected engorged mosquitoes along certain sections of east coast
Florida beach during turtle season (early May through mid September), his
results may have indicated that reptiles serve as an important vertebrate
host for salt marsh mosquitoes. This demonstrates the value of sampling a
wide variety of habitats over several active seasons before drawing general
conclusions from mosquito blood meal analysis. It also points out that even
when extensive sampling is done, as in Edman's study, pockets of unusual
blood feeding may be missed. While these areas of engorgement on unusual
vertebrate hosts may represent only a minor part of the overall species
specific feeding profile, they serve to document the extreme plasticity of
blood feeding patterns displayed by some mosquito species.
There is little question that salt marsh mosquitoes flying along ocean
dunes can and will respond opportunistically to the presence of sea turtles.
During our observation period, each female turtle fed hundreds of mos-
quitoes. It is unlikely that any other individual vertebrate host in the salt
marsh/sand dune habitat accounts for so many blood-fed mosquitoes in such
a short period of time.-J. F. DAY, Florida Medical Entomology Laboratory,
Vero Beach, FL 32960, USA, and G. A. CURTIS, Indian River Mosquito Con-
trol District, Vero Beach, FL 32960, USA.
A TECHNIQUE FOR SEPARATING MOLE CRICKETS FROM SOIL-
In order to determine mole cricket population densities and distributions in
the field, Williams and Shaw (1982. Florida Ent. 65: 192-4) developed an
efficient sampling technique which employs a tractor-powered soil corer.
This technique requires that the crickets then be separated from the large
quantities of soil. For this reason, a soil sifter was developed which was
portable, inexpensive, and yielded live crickets.
The sifter consists of a hopper made of 16 gauge galvanized steel (Fig.
1). The top perimeter has a 2.5 cm lip which supports the hopper on a re-
movable wooden frame made of 4.1 X 8.9 cm planks. A 0.63 X 0.63 cm mesh,
hardwarecloth sieve fits into the top and has 2 handles for easy removal. The
base perimeter of the hopper also has a 2.5 cm lip which provides a surface
for attaching a hatch. The hatch is secured by a piano hinge on one side and
by latches on the remaining sides. The latches are bolted to square pieces of
galvanized steel that are welded vertically from the edge of the lip to the
side of the hopper. The hatch is lined with a water-tight foam rubber gasket.
A hole in the front of the hopper is plugged with a rubber cork, which can be
removed to release excess water.
This technique relies simply on the principle that soils sink in water,
whereas crickets float or swim to the surface. The hopper is first filled with
ca. 20 liters of water, which prevents crickets from being buried by soil as
they fall through the sieve. The soil sample is spread on the sieve and forced
through with pressurized water from a garden hose. The sieve is scanned
for crickets and then removed from the hopper. At this stage the water
surface is typically covered with organic debris and foamy scum, which
360
September, 1983
Scientific Notes
361
+-FOAM RUBBER
Fig. 1. Portable soil sifter hopper and support frame for extracting mole
crickets from soil samples.
makes it difficult to see the crickets. A hand-held, straight-edged sieve (0.16
cm mesh) is used to skim the water. Its contents are washed into a 19-liter
bucket that has a 0.16 cm mesh screen for a bottom, and this is sprayed with
water until all the foam has passed through. The bucket sieve is then
placed inside another bucket filled with water, and the crickets are easily
spotted as they swim on the surface.
From 4 to 8 nineteen-liter samples can be processed per man h depending
on the amount of plant material and soil type. A blind test incorporating 10
nineteen-liter soil samples, 5 of which contained mole cricket nymphs in
various instars, was used to assay the accuracy of the technique (Table 1).
This test, and trials done in the field, showed that crickets are likely to be
missed in the hopper if they are crushed or die during core sampling or
sifting. In these instances, the lack of any movement by crickets among the
debris in the water makes it difficult to spot them, especially if they are 1st
instar nymphs.
An advantage of this technique is that crickets can be separated from
soil samples in situ provided there is a source of pressurized water. If the
source of water is mobile, then soil can be quickly returned to its point of
origin. The hopper is merely placed directly over the core hole and the soil
sample is deposited after sifting.
362 Florida Entomologist 66 (3) September, 1983
TABLE 1. NUMBER OF MOLE CRICKET NYMPHS INTRODUCED AND NUMBER
RECOVERED FROM FIVE 19-LITER SOIL SAMPLES USING A PORTABLE
SOIL SIFTER.
Soil sample
1 2 3 4 5 X
Crickets 10 7 6 5 3 -
introduced
Crickets 9 6 6 3 3 -
recovered
% 90 85.7 100 60 100 87.1
Materials for constructing the hopper and frame cost less than $60 and
both can be easily transported in a pickup truck or station wagon. The sifter
yields live crickets and could also be used to separate other soil dwelling
insects provided they are buoyant in water.
I thank T. J. Walker and W. T. Walker for their help and advice on this
project.-GARY N. FRITZ, Entomology-Nematology Dept., University of Flor-
ida, Gainesville, Florida 32611, USA. Present address: Dept. Entomology &
Plant Pathology, Box 3BE, New Mexico St. Univ., Las Cruces, NM 88001
USA.
LYRIFORM ORGANS IN THE CENTIPEDE SCUTIGERA (SCUTI-
GEROMORPHA: SCUTIGERIDAE)-A replica method for light micro-
scopy (K. T. Khalaf. 1980. Florida Ent. 63: 307-40) at 400 X magnification
was used to observe the cuticular pattern of the entire dorsum of the centi-
pede Scutigera sp.
The dorsal cuticle of the head and tergal plates possesses numerous
organs, each consisting of a striated tubercle and a slender companion
trichium (Fig. 1, 2). These appear to be similar to the lyriform organs of
the legs of some arachnids, which were illustrated by Barth (1976. Pages
445-73. In H. R. Hepburn, ed. Elsevier Scientific Publishing Co., New York).
Lyriform organs are composed of close and parallel slit sense organs.
Functionally, they serve as cuticular mechanoreceptors, stimulated by com-
pression of the slits. As far as I know, these organs have not previously been
reported from centipedes.
This investigation received support from the Academic Grant Fund of
Loyola University.-KAMEL T. KHALAF, Loyola University, New Orleans, LA
70118 USA.
Scientific Notes 363
Fig. 1, 2. Light micrographs of replicas of the integument of Scutigera
sp.: 1, head. 2, tergal plate. Total magnification X 1200.
FIREFLY CHROMOSOMES (PHOTINUS, PHOTURIS: LAMPYRIDAE
COLEOPTERA) -Over a one year period, Professor James Lloyd of the
University of Florida at Gainesville kindly collected, identified and sent to
us specimens of several species of fireflies. After a considerable amount of
preliminary difficulty and by employing protocols, somewhat modified, of
Lewis and Riles (1960. Drosophila Information Service 34: 118-9) and
Oster and Balaban (1963. Drosophila Information Service 37: 142-4), we
made the following observations with adult gonadal squashes:
Two Photinus macdermotti Lloyd males were found to have 19 chromo-
somes, the smallest being the X (Fig. 1). At diakinesis, one observes 10
bodies: one large ring bivalent which probably consists of a pair of
metacentrics with 2 terminal chiasmata; one smaller ring bivalent which
probably is a pair of acrocentrics with 2 terminal chiasmata; 7 pairs of
acrocentric bivalents each with a single terminalized chiasma; and a
univalent dot which is the X chromosome. The X chromosome was not seen
to be dividing. Stevens (1909. J. Exper. Zool. VI: 101-13), working with
species whose correct identity cannot be known with certainty, reported that
the males of both "Photinus consanguineus" and Photuris pennsylvanica
("Photinus pennsylvanicus" of Stevens; pennsylvanica has since been found
to be a complex of many species, Barber, 1951, Smithsonian Misc. Coll. 117, 1:
1-58) also have 19 chromosomes. Moreover, she reported that the X divides
Florida Entomologist 66 (3)
X
/
1 *t
1 io 6
Fig. 1. Composite drawing of 2 original photographs. Diakinesis of meiosis
of adult male Photinus macdermotti. "X" indicates X-chromosome.
during the first meiotic division in both species and that in "P. consan-
guineus" this occurs "very late, in a stage which is late anaphase or telophase
for the other chromosomes." In view of her observations and the fact that
we could not find any meiotic stages later than Metaphase I, we are not
prepared to say, at this time, when the X divides in P. macdermotti.
One Photuris congener Le Conte larva, sex unknown, had 18 chromosomes.
Several mitotic metaphase figures of spermatogonia (or oogonia) revealed a
pair of metacentric chromosomes, a pair of submetacentric chromosomes and
14 acrocentrics. We are now repeating and extending these observations to
other species in several genera. We thank Mrs. Alena Leff, Genetics Labora-
tory, Westchester County Medical Center, for aid with the initial histology.
-L. EHRMAN, State University of New York at Purchase, Purchase, NY
10577; M. WASSERMAN, Queens College, City University of New York, Flush-
ing, NY 11367 USA.
--^-- -^-- -- -^- -e *- - --
DAMAGE OF SLASH PINE FEMALE STROBILI BY REPRODUCTION
WEEVILS.-Reproduction weevils are commonly associated with branch
and stem debarking of pine seedlings planted in recently cutover pine-
forested lands. This note documents the occurrence of these weevils in the
crowns of larger trees and their damage to pine strobili.
On 23 February 1982, I observed 2 reproduction weevils feeding on the
female strobili of a 15 year old slash pine, Pinus elliottii Engelm., in a seed
orchard located in Munson, FL. The weevils were an adult $ pales weevil,
Hylobius pales (Herbst), and a 9 pitch-eating weevil, Pachylobius picivorus
(Germar). Both weevils were collected from the upper crown of the host
tree (ca. 11 m high).
A slash pine female strobilus and a flower stalk exhibited similar dam-
age: circular to linear excavation of plant tissue by the weevils (Fig. 1).
Pales weevils caused similar damages to Scots pine, Pinus sylvestris L., in
the Lake States (J. A. Corneil, pers. comm.). Contribution No. 542, Bureau
of Entomology, Division of Plant Industry, P. O. Box 1269, Gainesville, FL
32602, USA-W. N. DIxoN, Forest Entomologist, Division of Forestry,
Gainesville, FL 32602.
September, 1983
364
Scientific Notes
Fig. 1. (A) Slash pine strobilus damaged by Hylobius pales; (B) stalk
of slash pine strobilus damaged by Pachylobius picivorus (5.6 X). Photo
credit: J. Windsor (DPI Photo No. 7024444-13, -15).
365
Florida Entomologist 66 (3)
BOOK REVIEW
A CATALOGUE/CHECKLIST OF THE BUTTERFLIES OF AMERICA
NORTH OF MEXICO. 1981. Lee D. Miller and F. Martin Brown. Memoir
No. 2, The Lepidopterists' Society. 280 p. Members and subscribers, $10.00
cloth, $5.00 paper; non-members $17.00 cloth, $8.50 paper (U.S. currency).
This publication is the first catalogue of North American butterflies to
appear since Dyar's in 1902, and it must have been an herculean effort on the
part of the authors, considering the taxonomic changes that have ensued in
the interim. In the Introduction, Miller and Brown indicate that the
nomenclature used is based upon revisionary work through 1979, with some
1980 entries. For the most part, this is true, although some items were over-
looked. Other changes have occurred since the final copy went to press. In a
subsequent section of this review, a few of these matters are discussed.
In a fairly broad manner, the overall arrangement of families, genera and
species follows that proposed by dos Passos in his 1964 Checklist (Memoir
No. 1 of The Lepidopterists' Society), although considerable rearrangement
has taken place in some instances. The arrangement of genera within many
family groups is the reverse of the order proposed by dos Passos. The
Hesperioidea begin the treatment and the Danaidae end it, following the
Satyridae.
The nomenclature reflects trends in both western Europe and Japan with
the generic names of holarctic fauna following Higgins (1975. The Classifica-
tion of European Butterflies, Collins, London). Many names treated as sub-
genera by dos Passos have been elevated to full generic status by Miller and
Brown. A few old names have been resurrected, such as Basilarchia to re-
place Limenitis. At the species and subspecies levels, some taxa have been
removed from synonomy, while others have been sunk into synonomy. These
actions will undoubtedly provide grist for taxonomic mills for some time to
come.
Serious taxonomists will find this new catalogue extremely useful, al-
though I suspect that many will criticize the splitting of so many generic
names. Many non-specialists, however, will undoubtedly find the work per-
plexing, since it is a major departure from both the format and nomencla-
ture of the familiar dos Passos list.
The major difference between the dos Passos Checklist and this new opus
is the annotation. Memoir No. 2 is a full catalogue as well as a checklist.
Full citations for original descriptions are given, as well as location of type
specimens when known. In regard to the latter, however, all too often the
statements appear regarding location of types: "apparently in X museum"
or "type probably in X museum". This tends to be annoying when U.S.
museums are cited as possible repositories. Perhaps checking for type speci-
mens is not so easy as one would like to believe.
The original description citations end the frustration of trying to cross-
reference the dos Passos entries with Zoological Record, when frequently the
year citation in dos Passos was in error, or the name did not appear in
Zoological Record. References to clarifying footnotes appear throughout the
text. These footnotes comprise a 669-entry list at the end of the volume. The
index contains all of the scientific names that appear in the main text.
On first reading, the work seems free from excessive typographical
366
September, 1983
Book Review 367
errors, although some have been noted. Other errors and omissions are
treated in a subsequent section of this review.
The overall format and type selection provide a pleasing and very read-
able format. The binding of both the cloth and paper editions is excellent,
although the less durable spiral binding of Memoir No. 1 is perhaps more
convenient since the pages lie flat.
Although the authors relied primarily on published revisionary work,
they did not always follow such revisions exactly. In some instances, they
appear to have injected their own viewpoints. Such actions have produced a
few inconsistencies in the final treatment, some of which are noted in the
section-by-section discussion that follows.
HESPERIOIDEA-The taxa afranius (Erynnis) and albescens (Pyrgus) are
elevated to full species; harpalus and manitoba (Hesperia) are reduced to
subspecific rank under comma, following the trend in several recent pub-
lications. Only one North American subspecies of Carterocephalus palaemon
is recognized. This action should perhaps be reviewed in view of some of the
phenotypes in the western arctic. In Euphyes, ruricola has replaced vestris.
Although there is some evidence that Polites coras represents a South
American insect, this name has been retained over peckius. The species as-
signments in Agathymus follow Freeman's treatment (1969. J. Lepid. Soc.,
23 Suppl. 1: 1-59), rather than Roever's revision (in Howe ed., 1975. The
Butterflies of North America, Doubleday, New York). The authors might
do well to re-evaluate this action.
PAPILIONIDAE-The generic names used are indicative of recent trends in that
Papilio is used only for the machaon-complex species. Pterourus is used for
the tiger and spicebush swallowtail groups. Since there are some differences
between these 2 groups, as noted by Rothschild and Jordan (1906. Novitates
Zoologicae 13 (3) : 411-744), it is surprising that Euphoeades was not selected
for the tiger swallowtails, since a departure from Papilio had already been
made. The general species-subspecies assignments follow dos Passos with
oregonius as a separate species and dodi one of its subspecies. This varies
from some lists which place both oregonius and dodi as subspecies of bairdii.
Several problems appear in the treatment of Parnassius. In clodius,
shepardi is listed as a junior synonym of altaurus, an incorrect assignment.
The former is a large, relatively pale, red-spotted insect, while altaurus is a
medium-size, heavily maculated insect with yellow-orange spots. If shepardi
is to be sunk into synonomy, it should be under one of the northwestern
races such as claudianus or pseudogallatinus, as discussed by Ferris (1976.
J. Res. Lepid., 15(2): 65-74). Under P. phoebus, magnus and xanthus are
retained as valid subspecies, while montanulus and maximus are sunk into
synonomy. This is contrary to the reasons and actions expressed by Ferris
(1976. J. Res. Lepid., 15(1) : 1-22) and perhaps should be re-evaluated.
PIERIDAE-Generic nomenclature follows European trends. Colias harfordii
is retained as a full species, although ultraviolet studies indicate that it is a
subspecies of C. alexandra (Ferris, 1973. J. Lepid. Soc., 27(1): 57-73). A
recent revision proposed by Eitschberger in Germany (1981. Atalanta, 11(5) :
336-71) restricts Artogeia napi to the Old World, and suggests 4 species in
its place for the New World: venosa, oleracea, marginalis, angelika Eitsch-
berger (new species). This paper appeared after Miller and Brown was in
press, and the revisions proposed lack substantive supporting evidence.
Florida Entomologist 66 (3)
LYCAENIDAE-Generic nomenclature follows European trends; specific
nomenclature follows recent revisions with some exceptions. Euristrymon
is retained over Fixsenia as proposed by Clench (1978. J. Lepid. Soc., 32(4):
277-81). F. polingi organensis Ferris (1980. J. Lepid. Soc., 34(2) : 217-23) is
omitted. Incisalia mossii is retained as a subspecies of fotis, although treated
informally by many specialists as a separate species for the past decade.
Formal separation has been made by Fisher (in Ferris and Brown, 1981.
Butterflies of the Rocky Mountain States, U. Oklahoma Press, Norman).
Several new subspecies names proposed by Clench and published post-
humously (1981. Bull. Allyn Mus., 64: 1-31) for Mitoura siva and M.
spinetorum are not reflected in the Catalogue. The synonym boreale Lafon-
taine for S. borealis Lafontaine is omitted. Agriades aquilo and A. glandon
in North America have been replaced by the single species franklinii Curtis.
This is inconsistent with the Higgins and Riley treatment (1975. A Field
Guide to the Butterflies of Britain and Europe, Collins, London), in which
glandon is described as a montane species and aquilo as a low-elevation
coastal species. These 2 species also exhibit different larval host plant
preferences. On this basis franklinii is the North American counterpart of
aquilo. The W. H. Edwards name rustic is available to replace glandon for
the combinations r. rustica, r. lacustris, r. megalo, r. bryanti. The form
name caerulescenss" Ferris is incorrectly placed under Plebejus saepiolus
whitmeri; it should be under P. s. saepiolus. Icaricia icarioides helios (W. H.
Edwards) has been omitted.
RIODINIDAE-Little change from Memoir No. 1 other than the addition of
several names proposed by McAlpine in 1971.
LIBYTHEIDAE-NO basic changes; annotations.
HELICONIIDAE-Revisionary work included; annotations.
NYMPHALIDAE-The entries follow the dos Passos Checklist for the most
part, and the revisions thereto, but in reverse order. The revision of Speyeria
proposed in the Howe book is fortunately not followed. The taxon adiaste is
retained as a subspecies of egleis contrary to full species status as discussed
in Emmel and Emmel (1973. The Butterflies of Southern California, Nat.
Hist. Mus. of Los Angeles Co., Science Ser. 26: 1-148). In Speyeria zerene,
pfoutsi is treated as a valid subspecies rather than as junior synonym of
platina. The S. coronis gunderi-S. zerene controversy has not been resolved.
No comment appears under Boloria napaea concerning the revision proposed
by Crosson du Cormier (1977. Alexanor 10: 31-43) restricting napaea to the
Old World and elevating alaskensis to full species status.
A few items in the Melitaeini section require clarification. The combina-
tion Poladryas arachne nympha appears in the Catalogue, while Brown's
1966 paper cited in this entry shows the combination Poladryas minute
nympha. Poladryas minute (entry #591), now possibly extinct, appears
without subspecies. Thessalia fulvia is retained as a full species, while alma
is a subspecies of leanira. The Higgins revision of this group placed both
taxa as subspecies of leanira. The "first revisor" concept has been invoked
to replace Phyciodes campestris by P. pratensis, and 2 footnotes explain the
basis for this change. The Boisduval name pulchella has been omitted in the
Phyciodes tharos entries and does not appear in the index.
Asterocampa follows the single-species treatment of Reinthal (in Howe,
1975). Many specialists will take exception to this.
368
September, 1983
Book Review
Nymphalis vau-album watsoni (Hall) has been omitted.
SATYRIDAE-Numerous generic name changes occur based upon Miller's
continuing studies of the higher classification of North American Satyridae.
The Coenonympha entries generally follow the individual species arrange-
ment of the dos Passos list, rather than reflecting the tullia superspecies
approach of Ehrlich and Ehrlich (1961. How to Know the Butterflies, W. C.
Brown, Dubuque) and others. Some subspecific names in Erebia have been
elevated from synonomy without comment, such as E. epipsodea rhodia. Two
species of Erebia have been omitted entirely. These are erinnyn Warren
which was taken in the Yukon Territory in 1970, and E. inuitica Wyatt
(1966. Wien. ent. Gesl. Jg. 51: 93). The type locality of inuitica is "Endicott
Mts.", Alaska. Cercyonis oetus pallescens Emmel and Emmel (1971. Pan-
Pacific Ent. 47: 155-7) has been omitted as has also Coenonympha inornata
bottineauensis Chermock, Simmons and Chermock (in Knudson and Post,
July, 1963. Butterflies of Bottineau County, North Dakota Insects-Publ.
No. 2, Dept. of Agric. Ent., North Dakota State Univ.).
DANAIDAE-A few minor changes from the dos Passos Checklist appear.
Several other items merit comment. The type localities cited are as
originally published, but some are not presently in the original political
districts. For example Herschel Island (TL for Erebia young herscheli) is
now part of the Yukon Territory. The TL for Clossiana freija natazhati is
also now located in the Yukon Territory.
Species described or revised since the Catalogue went to press are:
Colias alexandra kluanensis Ferris (1981. Bull. Allyn Mus. 63: 1-12)
Callophrys (Xamia) xami texami Clench (1981. Bull. Allyn Mus. 64: 1-31)
Callophrys (Mitoura) siva chalcosiva Clench (op. cit.)
Callophrys (Mitoura) siva rhodope G. and S.-new status (op. cit.)
Callophrys (Mitoura) spinetorum ninus W. H. Edwards-elevated from
synonomy (op. cit.)
Specific errata noted are:
Footnote numbering not corrected in proof so that footnote "000" appears
on pages 60, 66, 116, 117, 121, 142, 190. The Roman numeral genus numbering
repeats itself on pages 91-92, thus offsetting the numbering in the remainder
of the text.
Entry #303a, the page citation under f. "pseudoamericus" should be 291, not
281.
Entry #311a, coloro is misspelled chloro and the name belongs with rudkini
(entry #305). The type was not destroyed; it is in the type collection in
The California Academy of Sciences.
Entry #343d, Euchloe hyanthis elsa. If one examines Beutenmiiller's original
description and illustration of this taxon, it appears that elsa is a varietal
form or aberration of creusa (entry #342) and not a subspecies of
hyantis.
Entry #636, interrogationis is misspelled interrogantionis.
Footnote #211, volume citation should be 33, not 34.
In a work of this magnitude, it is inevitable that omissions and errors
will occur. It is unfortunate that the manuscript as a whole was not circu-
lated to various specialists prior to publication. Many of the minor flaws and
problem areas would probably have been noted during this process. As noted
above, certain items require clarification, and one hopes that the authors
369
370 Florida Entomologist 66 (3) September, 1983
will publish a list of additions and corrections. Time will resolve many of the
nomenclatorial changes, many of which will appear radical to some users of
the Catalogue.
This Catalogue/Checklist is a vast improvement over existing literature
and contains a wealth of valuable information for taxonomic specialists and
amateur collectors alike. The literature citations and footnotes alone make
it well worth the modest cost. Memoir No. 2 should be on the bookshelf of
every lepidopterist interested in North American butterflies.-CLIFFORD D.
FERRIS, Bioengineering Program, University of Wyoming, Laramie, WY
82071; Research Associate, Florida State Collection of Arthropods, Division
of Plant Industry, Florida Department of Agriculture and Consumer Serv-
ices, Gainesville, FL 32602 USA.
North American Benthological Society 371
NORTH AMERICAN BENTHOLOGICAL SOCIETY
The North American Benthological Society is an international scientific
organization whose purpose is to promote better understanding of the biotic
communities of lake and stream bottoms and their role in aquatic ecosystems
by providing media for disseminating new investigation results, new interpre-
tations, and other benthological information to aquatic biologists and to the
scientific community at large. Membership is open to anyone interested in the
Society's purpose regardless of residence. Areas of major membership in-
terest include: ecology and life-histories of benthic biota; taxonomy or
systematics of aquatic biota; determination of pollution-tolerance ranges of
aquatic species; effect of water quality on distribution and abundance in the
benthic community; accelerated eutrophication; and methods of sampling and
measuring components of aquatic ecosystems.
NABS has a 3-day annual meeting with both verbal and poster presenta-
tions of scientific papers, symposia, workshops, and several social events. The
Society encourages graduate student participation at all levels and annually
presents awards for the best scientific paper presentations by graduate stu-
dents. As a service to its members and the scientific community at large, the
Society publishes the widely used Current and Selected Bibliography of
Benthic Biology, various nonperiodical symposia and workshop proceedings,
and the Bulletin of the North American Benthological Society (just start-
ing). Membership applications may be obtained from ELIZABETH B. RODGERS,
TVA Browns Ferry Biothermal Station, Box 2000, Decatur, AL 35602 USA.
372 Florida Entomologist 66 (3) September, 1983
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Sci-Mate operates on the Vector 3 or 4, the IBM Personal Computer, the
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