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(ISSN 0015-4040)
FLORIDA ENTOMOLOGIST
(An International Journal for the Americas)
Volume 74, No. 4 December, 1991
TABLE OF CONTENTS
Announcement 75th Annual Meeting ................................................ i
Research Reports
KRING, J. B., AND T. J. KRING-Variation in Body Shape, Number of Ocelli and
Number of Secondary Antennal Rhinaria on Wingless Males of Aphis sedi
(Homoptera: Aphididae) ............................ .............................. 487
WIRTH, W. W.-The Predaceous Midge Genus Allohelea Kieffer in the Western
Hemisphere (Diptera: Ceratopogonidae) ........................................... 491
WIRTH, W. W.-Forcipomyia bicolor and Related Species of the Subgenus
Lepidohelea in Brazil (Diptera: Ceratopogonidae) ............................. 506
SKELLEY, P. E., AND R. E. WOODRUFF-Five New Species ofAphodius (Coleop-
tera: Scarabaeidae) From Florida Pocket Gopher Burrows .................. 517
THOMAS, M. C., AND S. B. PECK-Survey of Insects of South Florida and the
Florida Keys: Flat Bark Beetles (Coleoptera: Cucujidae [sensu. lat.]
[Laemophloedae, Passandridae, Silvanidae]) .................................... 536
ROACH S. H.-Natural Plant Materials as Overwintering Sites for Arthropods in
the Coastal Plain of South Carolina ............................................... 543
WALKER, T. J., AND S. A. WINRITER-Hosts of a Phonotactic Parasitoid and
Levels of Parasitism (Diptera: Tachinidae: Ormia ochracea) ................ 554
GALLIART, P. L., AND K. C. SHAW--Effect of Intermale Distance and Female
Presence on the Nature of Chorusing by Paired Amblycorypha parvipennis
(Orthoptera: Tettigoniidae) Males ............................................... 559
GREANY, P. D., R. E. MCDONALD, W. J. SCHROEDER, AND P. E. SHAW-
Improvement in Efficacy of Gibberellic Acid Treatments in Reducing
Susceptibility of Grapefruit to Attack by Caribbean Fruit Fly ............. 570
Scientific Notes
PETITT, F. L.-Effect of Photoperiod on Larval Emergence and Adult
Eclosion Rhythms in Liriomyza sativae (Diptera: Agromyzidae) ... 581
DOWNING, A. S., C. G. ERICKSON, AND M. J. KRAUS-Field Evaluation
of Entomopathogenic Nematodes Against Citrus Root Weevils (Col-
eoptera: Curculionidae) in Florida Citrus ............................... 584
IGNOFFO, C. M., AND C. GARCIA--Conidial Germination of Two Biotypes
of Nomuraea rileyi ............................ .......... ... 587
MINNO, M. C.-Outbreak of Variable Oakleaf Caterpillar, Lochmaeus man-
teo (Lepidoptera: Notodontidae) at the Katharine Ordway Preserve-
Swisher Memorial Sanctuary, Putnam County, Florida ............. 589
ALI, A., AND M. L. KOK-YOKOMI-Preliminary Population Assessment
of Psychoda alternate (Diptera: Psychodidae) in Soil Irrigated with
Wastewater for Turf Cultivation ................. .. ......... 591
Continued on Back Cover
Published by The Florida Entomological Society
FLORIDA ENTOMOLOGICAL SOCIETY
OFFICERS FOR 1990-91
President ................................................................................ J. L. Knapp
President-E lect .................... ............................................... D F. W illiam s
Vice-President .............. ..................................................... J. E. Pela
Secretary ...................................... ................... ......... ........ D. G. Hall
Treasurer .................... .............................................. A. C. Knapp
Other Members of the Executive Committee
J. F. Price J. E. Pefa J. R. Cassani
J. R. McLaughlin O. Liburo F. Oi
PUBLICATIONS COMMITTEE
J. R. McLaughlin, USDA/ARS, Gainesville, FL ....................................... Editor
Associate Editors
Agricultural, Extension, & Regulatory Entomology
James R. Brown-Disease Vector Ecology & Control Center, NAS, Jacksonville, FL
Richard K. Jansson-Tropical Research & Education Center, Homestead, FL
Michael G. Waldvogel-North Carolina State University, Raleigh, NC
Apiculture
Stephen B. Bambara-North Carolina State University, Raleigh, NC
Biological Control & Pathology
Ronald M. Weseloh-Connecticut Agricultural Experiment Sta., New Haven, CT
Book Reviews
J. Howard Frank-University of Florida, Gainesville
Chemical Ecology, Physiology, Biochemistry
Louis B. Bjostad-Colorado State University, Fort Collins, CO
Ecology & Behavior
Theodore E. Burk-Creighton University, Omaha, NE
John H. Brower-Stored Product Insects Research Laboratory, Savannah, GA
Forum & Symposia
Carl S. Barfield-University of Florida, Gainesville
Genetics & Molecular Biology
Sudhir K. Narang-Bioscience Research Laboratory, Fargo, ND
Medical & Veterinary Entomology
Arshad Ali-Central Florida Research & Education Center, Sanford, FL
Resumen
J. E. Pena-Tropical Research & Education Center, Homestead, FL
Systematics, Morphology, and Evolution
Michael D. Hubbard-Florida A&M University, Tallahassee
Howard V. Weems, Jr.-Florida State Collection of Arthropods, Gainesville
Willis W. Wirth-Florida State Collection of Arthropods
Business Manager .............................................. ...................... A. C. Knapp
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This issue mailed December 20, 1991
ANNOUNCEMENT OF THE 75TH ANNUAL MEETING
FLORIDA ENTOMOLOGICAL SOCIETY
The 75th annual meeting of the Florida Entomological Society will be held August
10 13, 1992 at the Indian River Plantation Resort and Conference Center, 555 N.E.
Ocean Boulevard, Hutchinson Island, Stuart, Florida 34996; telephone (407) 225-3700;
Fax (407) 225-0003. Registration forms and information will be mailed to members and
will appear in the Newsletter.
SUBMISSION OF PAPERS
The deadline for submission of papers and posters for the 75th annual meeting of
the Florida Entomological Society will be May 15, 1992. The meeting format will be
similar to those in the past with eight minutes allotted for presentation of oral papers
(with two minutes for discussion) and separate sessions for members who elect to pres-
ent a Poster Exhibit. There will be student paper and poster sessions with awards as
in previous years. Students participating in the judged sessions must be members of
the Florida Entomological Society and registered for the meeting.
Jorge E. Pefia, Chairman
Program Committee, FES
University of Florida
Tropical Research and Education Center
18905 S.W. 280 Street
Homestead, Florida 33031
(305) 246-7048 or 246-6340
Kring & Kring: Wingless Males of Aphis sedi
VARIATION IN BODY SHAPE, NUMBER OF OCELLI
AND NUMBER OF SECONDARY ANTENNAL RHINARIA
OF WINGLESS MALES OF APHIS SEDI
(HOMOPTERA: APHIDIDAE)
JAMES B. KRING
Gulf Coast Research and Education Center
University of Florida, FL 34203
Deceased.
TIMOTHY J. KRING
Department of Entomology
University of Arkansas
Fayetteville, AR 72701
ABSTRACT
In a collection of sexual forms of Aphis sedi Kaltenbach, wingless males were found
with alate-like bodies and others with apterous-like bodies. These individuals varied in
the presence or absence of ocelli and in the number of secondary rhinaria on the antenna.
RESUME
Al colectar las formas sexuales de Aphis sedi Kaltenbach, se encontraron machos
sin alas con cuerpo similar a el de los machos alados y otros con cuerpo similar a el de
los machos sin alas. Se diferencio en estos especimenes la presencia y la ausencia del
oceli y el numero de rhinaria secundaria en la antena.
In the sub-family Aphidinae, most male aphids are alate and most oviparous females
are apterous (Hille Ris Lambers 1966). Alate morphs possess 3 ocelli, both primary and
secondary compound eyes and secondary rhinaria on the antennae. Apterous individuals
lack ocelli, may lack secondary compound eyes and may also lack or have fewer secondary
antennal rhinaria (Hille Ris Lambers 1966, Miyazaki 1987). Secondary rhinaria are
important in mate location (in males) and host plant location (in females) (Anderson &
Bromley 1987). In many of these species, the alate male is born on one plant species
and must locate oviparous females born on another. In a smaller number of species,
both sexes lack wings and are produced on the same plant. Males of the Aphidinae that
lack wings often are referred to as intermediate or apterous males. These wingless
males sometimes have the body shape of alate aphids and are not intermediate in male
structures. True apterous males differ in body shape from alate aphids in that they do
not have a distinct thorax. Neither of these terms is satisfactory.
Do these wingless males, which cannot make effective use of host or mate-locating
organs, lack or have fewer ocelli or secondary antennal rhinaria? The answer to this
question is not in descriptions of these aphid species in the literature. However, it could
be expected that wingless males might lack ocelli, since in the species Macrosiphoniella
sanborni Gillette, the determination and differentiation of ocelli of the alate viviparous
female is related to the development of wings and wing muscles (Kitzmiller 1950).
Aphis sedi Kaltenbach, a species rare in New England, occurs year-round on Sedum
sp. or Sempervivum sp. At this latitude (New Haven, CT, USA) apterous oviparous
females and wingless males can appear on infested plants in October. These mate and
488 Florida Entomologist 74(4) December, 1991
the females deposit the overwintering eggs. There is no known alternate host (Kring
1955).
Here we describe the occurrence of ocelli and secondary antennal rhinaria on wingless
males of A. sedi and relate this information to the characters of other morphs of this
species (Kring 1955, Theobald 1927).
METHODS AND MATERIALS
The aphids described here were collected from Sedum sp. in October in Branford,
CT. Over 100 adults were examined. Aphids were killed in 70% ethyl alcohol, macerated
for 24 hours in cold 10% NaOH and then gradually dehydrated with increasing concen-
trations of ethyl alcohol. The specimens were cleared in clove oil. The clove oil was
gradually mixed with the mounting medium diaphane to reduce shrinkage. They were
preserved in pure diaphane on glass slides under cover glasses. These aphids were
examined under a compound microscope, and the number of ocelli and secondary antennal
rhinaria were recorded.
RESULTS AND DISCUSSION
The general body characteristics of one of the wingless males of A. sedi (Fig. 1A)
are those of an alate aphid. There is a distinct prothorax and a fused meso- and
metathorax. The thorax of this A. sedi male is about the same width as the head and
is distinct from both the head and the abdomen. In the apterous aphid (Fig. 1B), it is
difficult to distinguish these sections of the body and there is a rapid increase in the
width of the body posterior to the metathoracic legs. The meso- and the metathoracic
tergal plates in these wingless males do not appear to be as distinct, nor as divided, as
those of the alate aphid.
Among the nine male individuals examined from this collection some males lacked
ocelli and others possessed one, two or three. Examples were selected to illustrate the
observed variation (Table 1). When only one ocellus was present, it was the medial
ocellus. When two were present, they were the medial and the left dorsal ocellus. If all
three were present the right ocellus was sometimes reduced (Fig. 1A). Some individuals
lacking ocelli had an indentation where the medial ocellus should be located. If all three
were absent the body was apterous shaped. Individuals having a reduced number of
ocelli also showed differences in body shape. The thorax of these insects were indistinct
and wing muscles were absent.
Wingless A. sedi males and alate vivaparous females possess both primary and
secondary rhinaria on the antennae. Apterous oviparous and viviparous females have
only primary rhinaria on the antennae, one on antennal segment V and one on antennal
segment VI (Kring 1955). Alate viviparous females possess these primary rhinaria, and
in addition 5 to 8 secondary rhinaria per antenna, usually all located on segment III
(Theobald 1927). These wingless males have the primary rhinaria and also 19 to 36
secondary rhinaria per antenna, on segments III, IV and V (Table 1). The antenna of
the wingless male is longer than that of the oviparous or viviparous female. Those males
lacking or having a reduced number of ocelli also have fewer secondary rhinaria on their
antennae (Table 1).
The information presented here describes some of the polymorphism that can occur
among males in A. sedi. This is the first report that shows reduction in the number of
ocelli on a male aphid and relates this loss to diminished numbers of secondary rhinaria
on the antennae and changes in body shape.
Kring & Kring: Wingless Males of Aphis sedi
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December, 1991
Florida Entomologist 74(4)
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Wirth: Western Hemisphere Allohelea
ACKNOWLEDGMENTS
We thank Dr. Dave Schuster, GCREC, University of Florida and anonymous review-
ers for their suggestions. This is Florida Agricultural Experiment Station Journal Series
No. R-01140. Please address reprint requests to second author.
REFERENCES CITED
ANDERSON, M., AND A. K. BROMLEY. 1987. Sensory system, in Aphids: their biology,
natural enemies and control. Vol. 2A: 153-162. Elsevier Sci. Pub. Amsterdam.
The Netherlands.
Hille Ris Lambers, D. 1966. Polymorphism in Aphididae. Ann. Rev. of Entomol. 11:
47-78.
KITZMILLER, J. B. 1950. The time interval between determination and differentiation
of wings, ocelli and wing muscles in the aphid, Macrosiphon sanborni (Gillette).
Amer. Nat. 84: 23-50.
KRING, J. B. 1955. Biological separation of Aphis gossypii Glover and Aphis sedi
Kaltenbach. Ann. Entomol. Soc. Amer. 48: 442-444.
MIYAZAKI, M. 1987. Forms and morphs of aphids. in Aphids: their biology, natural
enemies and control. Vol. 2A: 27-47. Elsevier Sci. Pub. Amsterdam. The Nether-
lands.
THEOBOLD, F. V. 1927. The plant lice or Aphididae of Great Britain. Vol. 2. Headley
Brothers. London. 411 pp.
THE PREDACEOUS MIDGE GENUS ALLOHELEA KIEFFER
IN THE WESTERN HEMISPHERE (DIPTERA:
CERATOPOGONIDAE)
WILLIS W. WIRTH
Research Associate, Florida State Collection of Arthropods, and
Cooperating Scientist, Systematic Entomology Laboratory, USDA
1304 NW 94th St., Gainesville, Florida 32606
ABSTRACT
Redescriptions and figures are presented for the two previously known American
species of Allohelea Kieffer, nebulosa (Coquillett) and johannseni (Wirth), and distribu-
tion records and misidentifications are corrected. Five new species are described: bottim-
eri from Texas, distortifemur, pedicellata and weemsi from Florida, and neotropica
from the Caribbean, Central America and northern South America. A revised diagnosis
is given for the genus Allohelea, and a key is presented for the western hemisphere
species.
RESUME
Se presentan nuevas descripciones y figures de 2 species americanas las cuales eran
conocidas previamente como Allohelea Keiffer nebulosa (Coquillett) y johannseni
(Wirth), y se corrige la information sobre su distribution e identificaciones previas. Se
described 5 species nuevas: baltimeri de Texas, distortifemur, pedicellata y weemsi
de Florida y neotropica del Caribe, Centromamerica y la parte norte de Sur America.
Florida Entomologist 74(4)
Se hace una revision de la diagonsis para el genero Allohelea, y se present una clave
para las species del hemisferio occidental.
Study of extensive Florida collections of the predaceous midges of the genus Allohelea
Kieffer has revealed the presence of three distinctive and previously undescribed species
and disclosed confused identifications in the two previously known Nearctic species, A.
nebulosa (Coquillett) and A. johannseni (Wirth). This paper will attempt to correct
previous errors and will present descriptions of the three new Florida species as well
as two additional species from Texas and the Neotropical Region that were previously
misdetermined. A revised diagnosis is given for the genus Allohelea, and a key is
presented for the Western Hemisphere species.
Types of the new species are deposited in the National Museum of Natural History,
Smithsonian Institution, Washington, D.C. Paratypes, when available, will be deposited
in the Florida State Collection of Arthropods, Gainesville; California Academy of Sci-
ences, San Francisco; British Museum (Natural History), London; and Museum National
d'Histoire Naturelle in Paris.
Terminology of taxonomic characters follows that of Downes & Wirth (1981) and
Wirth & Grogan (1988). Measurements are given in microns unless otherwise specified.
Genus Allohelea Kieffer
(Fig. 1)
Allohelea Kieffer, 1917: 364. Type-species, Sphaeromias pulchripennis Kieffer, by orig-
inal designation.
REFERENCES: Wirth, 1953: 135 (Monohelea tesselata Group; America); Wirth
& Williams, 1964: 304 (Monohelea tesselata Group; North American species); Lane &
Wirth, 1964: 213 (Monohelea tesselata Group; Neotropical species); Wirth & Grogan,
1988: 14 (generic status; diagnosis; world list of species).
Diagnosis. Moderately large, stout biting midges; wing length 1.2-2.0 mm.
Female antenna (Figs. 2, 8) moderately long; male antenna with plume, segments 3-12
fused. Palpus (Figs. 3, 9) with third segment moderately swollen, with small round
sensory pit. Female mandible with about eight strong teeth. Wing (Figs. 1, 5) with
three large dark anterior patches and irregular posterior infuscated areas more or less
of same intensity; two well-formed radial cells; costal ratio 0.72-0.84. Femur and tibia
of hind leg (Fig. 4) swollen and shining black. Hind tarsus (Figs. 4, 10) bearing very
strong ventral spines at apices of proximal 2-3 tarsomeres; fourth tarsomeres of fore
and mid legs cylindrical, of hind leg elongated; claws of fore and mid legs of female
short, bent basally, straight distally, with basal internal and external teeth and often
bifid tips; in male small, equal and simple, tips bifid; hind claw (Figs. 4, 10) of both
sexes single, long, usually with short basal tooth. Female tenth sternum (Fig. 7) with
4-6 pairs of setae; two more or less equal sized spermathecae (Figs. 15-19). Male aedeagus
(Fig. 6) without basal loop, the elongate, tapering distal process borne in a notch in the
quadrate basal sclerite; parameres (Fig. 6) joined mesad near bases, posterior portion
simple and tapering to pointed tip directed more or less dorsocaudad over distal portion
of ninth tergum.
Distribution. Worldwide, mainly Oriental; 38 species.
December, 1991
Wirth: Western Hemisphere Allohelea
493
Fig. 1. Allohelea nebulosa (Coquillett), female.
Key to the American Species of Allohelea
1. Females ..................................................... ............................. 2
- Males ............................................................................................. 8
2. Hind femur modified at base with slender subcylindrical pedicel (Fig. 11) or
ventral excavation and knob (Fig. 10) ..................................... .......... 3
- Hind femur without pedicel or excavation and knob (Figs. 13-15) .............. 4
3. Hind femur with two ventral constrictions separated by a knoblike swelling
on basal half (Fig. 10) ................................................ distortifemur n. sp.
- Hind femur with abrupt subcylindrical constriction on basal third (Fig. 11)
.................................................................................. pedicellata n. sp.
Florida Entomologist 74(4)
4. Hind femur gradually tapered to slender base (Fig. 13); (larger species, wing
length 1.57-1.74 mm) ................................................ nebulosa (Coquillett)
- Hind femur uniformly stout (Figs. 14, 15) or with short, abrupt, basal con-
striction (Fig. 11) ................................................ ...................... 5
5. Smaller species, wing length 0.99-1.39 mm; hind femur not greatly swollen,
slightly tapering to base; spermathecae less than 102 microns long ........... 6
- Larger species, wing length 1.65 mm; hind femur with short abrupt basal
constriction, greatly and evenly swollen to tip (Fig. 11); spermathecae 115-
122 microns long ................................................................. weemsi n. sp.
6. Hind femur with yellow knee spot at tip, wing length 0.99 mm ..............
................................................................................ neotropica n. sp.
- Hind femur dark to tip; wing length 1.19-1.39 mm .................................. 7
7. Wing length 1.25-1.39 mm ........................................... johannseni (Wirth)
- Wing length 1.19 mm ...................................................... bottimeri n. sp.
8(1). Hind femur with yellow knee spot; hind claw without basal tooth; small
species, wing length 1.09 mm ........................................ neotropica n. sp.
- Hind femur dark to tip; hind claw with basal tooth ................................. 9
9. Dististyle with stout, blunt tip; distal sclerite of aedeagus as broad as long
(Fig. 22); small species, wing length 1.11 mm ...................... bottimeri n. sp.
- Dististyle tapered to slender tip; distal sclerite of aedeagus much longer
than broad (Figs. 23, 25, 26); wing length 1.17-1.58 mm ......................... 10
10. Aedeagus with large, sagittate distal expansion (Fig. 26) ........... weemsi n. sp.
- Aedeagus tapering distally to slender tip (Figs. 6, 23, 26) ....................... 11
11. Dististyle short, 0.50 length of basistyle, stout at base and tapering to sharp-
pointed tip (Fig. 25); wing length 1.44 mm .................... distortifemur n. sp.
- Dististyle longer, 0.58-0.75 length of basistyle, more slender proximally.
(Figs. 6, 23) .................................................. ............................. 12
12. Dististyle with more slender tip (Fig. 23); wing length 1.58 mm .............
............................................................................. nebulosa (Coquillett)
- Dististyle with stouter tip (Fig. 6); wing length 1.17 mm ... johannseni (Wirth)
Allohelea nebulosa (Coquillett)
(Figs. 1, 13, 21, 23)
Ceratopogon nebulosus Coquillett, 1901: 606 (male; New Jersey).
Ceratolophus nebulosus (Coquillett); Kieffer, 1906: 60 (combination).
Johannseniella nebulosa (Coquillett); Malloch, 1914: 226 (combination).
Hartomyia nebulosa (Coquillett); Malloch, 1915: 340 (combination; Indiana; notes).
Monohelea nebulosa (Coquillett); Kieffer, 1917: 312 (combination); Johannsen, 1943: 781
(distribution); Wirth, 1953: 151 (redescribed; figs.; distribution); Wirth & Williams,
1964: 304 (notes; figs.; distribution); Wirth & Grogan, 1981: 44 (diagnosis; figs.;
Potomac Valley records).
Allohelea nebulosa (Coquillett); Wirth & Grogan, 1988: 15 (combination).
Female (Fig. 1). Wing length 1.57-1.74 mm; breadth 0.65 mm; costal ratio 0.80.
Head: Dull dark brown with a few pruinose gray spots on vertex; antenna light
brown, distal segments not darker; lengths of flagellar segments 72-43-47-50-54-58-58-58-
82-82-82-82-94; antennal ratio 0.97. Palpus light brown, third segment variably yellowish;
lengths of segments 18-43-72-50-72; palpal ratio 2.5. Mandible with eight teeth.
Thorax: Dark brown; mesonotum pruinose gray, with scattered small dark brown
spots at bases of setae, these spots more or less confluent, especially on posterior portion
of mesonotum; scutellum with middle third variably yellowish. legs shining dark brown,
December, 1991
Wirth: Western Hemisphere Allohelea
ci)ELZAZ-DKDDCCChJ
- ---i
I ----
Figs. 2-7. Alloheleajohannseni: 2-5, 7, female; 6, male: 2, antenna; 3, palpus; 4, hind
leg; 5 wing; 6, 7, genitalia.
tarsi stramineous. Hind leg with femur and tibia greatly thickened, femur (Fig. 13)
variably narrowed at base, somewhat petiolate; lengths from femur to T5, 980-860-340-
180-94-122-122; claw 165, basal tooth 58; hind basitarsus with basal and distal spine,
second tarsomere with two distal spines, third with one distal spine. Wing (Fig. 1)
grayish hyaline; a quadrate black patch across basal cell at half its length; two broad,
irregular, more or less interconnected transverse bands across wing, the first at first
496 Florida Entomologist 74(4) December, 1991
radial cell, second at apex of second; cell R5 thus with a dark, a light, a dark, and a
light band, all of subequal breadth; first radial cell 0.4 length of second. Halter pale yellow.
Abdomen: Pruinose dark brown, variably paler to yellowish at base; cerci yellowish.
Spermathecae (fig. 21) oval with short slender necks; measuring 102 x 65 and 87 x 61,
necks 9.
Male. Wing length 1.55-1.70 mm; breadth 0.45 mm; costal ratio 0.71.
Similar to female with usual sexual differences. Antenna with segments 3-12 fused,
plume yellowish at base; lengths of flagellar segments 150-47-47-51-51-51-51-51-47-72-
122-108-115. Hind leg with femora and tibiae not quite as swollen as in female; lengths
from femur to T5, 970-900-290-172-88-122-115, claw 144, basal tooth 29.
Genitalia (Fig. 23): Posterior margin of ninth sternum transversely abutting base of
aedeagus; ninth tergum moderately short and greatly tapering distally, sides subparallel
on distal half, apicolateral processes short and rounded. Basistyle stout and slightly
curved; dististyle slender and curved, 0.75 length of basistyle. Aedeagus with basal
sclerite about as broad as long, anterior margin contiguous with ninth sternum, posterior
margin deeply emarginate, the tapering rodlike distal sclerite fitting proximally into
the notch, swollen at base and flanked by a membranous sheath variably appressed to
it or parted along ventral line in low irregular flaps. Parameres connected by a difficult-to-
see transverse bridge near bases, with slender anterior processes; each paramere with
posterior portion narrowed and curved laterad and then dorsomesad to a bladelike tip
tucked dorsally around ninth tergum.
Distribution. Nearctic; Wisconsin to Massachusetts, south to Arkansas and Florida.
Type. Holotype male, Riverton, New Jersey, C. W. Johnson (Type no. 5479, USNM).
Specimens Examined. ALABAMA: Baldwin Co., Gulf Shores, Gulf State Park, 28-
31.v.1978, J.I. Glick, 2 males; Bog off Co. 47, 2 km w 1-65, in Sarracenia purpurea,
12.v.1978, J.I. Glick, 1 male. Cleburne Co., Cheaha Mtn., Cheaha State Park, 29-
31.v.1978, J.I. Glick, 1 male, 1 female. Lee Co., Chewacla State Park, 20-21.iv.1977,
J.I. Glick, 2 males. Mobile Co., Dauphin Island, Heron Bayou, 14.v.1977, J.I. Glick &
G. R. Mullen, 1 female; Ft. Gaines Campground, 13.v.1977, Glick & Mullen, 1 male.
ARKANSAS: Drew Co., Monticello, 13.v.1969, G. Hatley, 1 female. Pike Co., 12.vi.1936,
W. F. Turner, 1 female. FLORIDA: Alachua Co., Gainesville, Chantilly Acres, v.1967,
F. S. Blanton, light trap, 2 males, 7 females; 23.iv.1970, W. W. Wirth, malaise trap, 1
female; Hawthorne, 27.iv.1968, F. S. Blanton, light trap, 1 female; High Springs,
20.iv.1954, M. Nelson, light trap, 1 female. Baker Co., Glen St. Mary, v.1971, F. S.
Blanton, UV light trap, 2 males; Olustee, vii.1971, F.S. Blanton, light trap, 1 male.
Gulf Co., St. Joseph State Park, 1.v.1970, W. W. Wirth, light trap, 17 males, 3 females.
Jefferson Co., Monticello, v.1969, W. H. Whitcomb, UV light trap, 3 males, 3 females.
Liberty Co., Torreya State Park, 15.iv.1957, 27.iv.1958, F. S. Blanton, 3 males, 16
females; 15.v.1971, G. B. Fairchild, UV light trap, 2 males; 20.v.1966, H. V. Weems,
Jr., 4 males, 12 females; 22.iv.1967, W. W. Wirth, light trap, 2 males, 7 females. Marion
Co., Juniper Springs State Park, 28.iv. 1970, W. W. Wirth, light trap, 2 males, 2 females.
Orange Co., Rock Springs, 21.iv.1970, W. W. Wirth, UV light trap, 1 male, 2 females.
Santa Rosa Co., Blackwater River Biol. Sta., 21.v.1974, G. B. Fairchild, UV light trap,
1 female; Jay, v.1962, T. W. Boyd, light trap, 2 females. Wakulla Co., Ochlockonee
River State Park, 29.iv.1970, W. W. Wirth, light trap, 1 male. MARYLAND: Prince
Georges Co., Patuxent Wildlife Res. Ctr., vi.1976, W.L. Grogan, Jr., malaise trap, 1
male. Wicomico Co., Salisbury, vi-viii.1979, W. L. Grogan, Jr., malaise trap, 2 males,
4 females. Worcester Co., Snow Hill, 9.vii.1966, W. H Anderson, light trap, 5 males,
9 females. MASSACHUSETTS: Middlesex Co., Bedford, 21.vii.1961, W. W. Wirth,
swept in swamp, 4 females. NORTH CAROLINA: Jackson Co., Dulaney Bog, 10 km
S Cashiers, 21.vi.1986, W. W. Wirth, malaise trap, 2 females. Washington Co., Pettigrew
Wirth: Western Hemisphere Allohelea
8
~~ccz
o~-~i~=C9
15 3 1
15 19
Figs. 8-19. Allohelea spp., female; 8-10, 16, distortifemur; 11, weemsi; 12, 20, pedicel-
lata; 13, 21, nebulosa; 14, 19, johannseni; 15, 17, neotropica; 18, bottimeri; 8, antenna;
9, palpus; 10, hind leg; 11-15, hind femur; 16-21 spermathecae.
State Park, 3.vi.1959, D. A. Young, 1 male. WISCONSIN: Shawano Co., Shawano,
vi.1968, R. Habeck, 1 male.
Discussion. Allohelea nebulosa is difficult to distinguish from A. johannseni (Wirth),
but averages larger in size (wing length 1.63 vs. 1.30), the mesonotal pattern pruinose
gray with small punctiform spots vs dark brown with large pruinose gray areas, the
distal antennal segments are longer and more slender, the female hind femur is more
slender and tapered toward the base, and the male dististyle is longer and much more
slender distally.
This species has been confused in the past literature with other species; Wirth &
Williams (1964) figured the male aedeagus from Texas specimens that are herein de-
scribed as A. bottimeri n. sp.
: "21
Q
Florida Entomologist 74(4)
Allohelea johannseni (Wirth)
(Figs. 2-7, 14, 19)
Monohelea johannseni Wirth, 1953: 153 (male, female; Virginia; figs.); Wirth &
Williams, 1964: 305 (notes; distribution; figs.); Wirth & Grogan, 1981: 39 (diagnosis;
Potomac Valley records; figs.); Graves & Graves, 1985: 88 (North Carolina; reared from
Boletus fungi).
Allohelea johannseni (Wirth); Wirth & Grogan, 1988: 15 (combination).
Female. Wing length 1.06-1.30 mm; breadth 0.51 mm; costal ratio 0.78. Closely
resembling Allohelea nebulosa, but smaller; mesonotum dark brown with large, irregu-
lar, pruinose gray areas; scutellum entirely blackish and with four long and several
shorter marginal setae. Wing pattern (Fig. 5) as in A. nebulosa, but markings behind
vein M1 much fainter than those on anterior part of wing. Head: Antenna (Fig. 2) with
lengths of flagellar segments 54-43-43-47-47-47-50-50-72-72-72-72-93; antennal ratio 1.00.
Palpus (Fig. 3) with lengths of segments 24-32-57-36-72; palpal ratio 2.0. Mandible with
eight teeth.
Thorax: Scutum subshining black with reduced pattern of pruinose gray areas; scutel-
lum black. Hind leg (Figs. 4, 14) with femur evenly swollen to base, without apical
yellow knee spot; lengths of segments from femur to T5, 790-720-260-144-94-108-108;
claw 166, basal tooth 53.
Abdomen: Genital segments as in Fig. 7; spermathecae (Fig. 19) 75 x 54 and 61 x
47, with necks 11.
Male. Wing length 1.05-1.20 mm; breadth 0.48 mm; costal ratio 0.63. As in female
with usual sexual differences. Antenna with lengths of flagellar segments 122-43-43-43-
36-36-36-36-36-54-108-90-100. Hind leg slightly more slender than in female; lengths from
femur to T5, 720-680-214-144-65-86-94; claw 144, basal tooth 36.
Genitalia (Fig. 6): As in A. nebulosa, but dististyle with more bluntly pointed tip.
Aedeagus without convex basal swelling on distal sclerite, the lateral flanking membrane
poorly developed.
Distribution. Nearctic; Ontario to New Brunswick, south to Kansas and Florida.
Types. Holotype female, Falls Church, Fairfax Co., Virginia, 8.vii.1950; W. W.
Wirth (type no. 61095, USNM). Allotype male, Innerarity Point, Escambia Co., Florida,
29.iv.1949, Rathert, light trap. Paratypes, Virginia, Florida, Alabama, Michigan, Iowa,
4 males, 9 females.
Specimens Examined. ALABAMA: Baldwin Co., Gulf Shores, Gulf State Park,
18.iv.1969, G. M. Stokes, light trap, 1 male; 28-31.v.1978, J. I. Glick, 2 males. DeKalb
Co., DeSoto State Park, 30.v-5.vi.1978, J. I. Glick, 1 male. Macon Co., Tuskegee Nat.
Forest, sphagnum bog, 15.iv.1978, J. I. Glick, 1 female. Mobile Co., Dauphin Island,
Ft. Gaines Campground, 13.v.1977, J. I. Glick, 3 females. FLORIDA: Alachua Co.,
Gainesville, Chantilly Acres, v-vi.1967, F. S. Blanton, light trap, 8 males, 13 females.
Baker Co., Glen St. Mary, v.1971, F. S. Blanton, UV light trap, 2 females. Bay Co.,
Beacon Hill, 8.iv.1971, F. S. Blanton, UV light trap, 1 male, 55 females. Escambia Co.,
Bratt, vi.1968, F. S. & A. J. Blanton, light trap, 2 females; Innerarity Point, 29.iv.1949,
Rathert, light trap, 5 males, 6 females; Molina, 23.viii.1969, F. S. Blanton, light trap,
1 male. Franklin Co., Wright Lake nr Sumatra, iv.1973, G. B. Fairchild, UV light trap,
3 males, 2 females. Gulf Co., St. Joseph State Park, 1.v.1970, W. W. Wirth, light trap,
1 female; Wewahitchka, v.1961, St. Bd. Health, light trap, 1 female. Highlands Co.,
Lake Placid, Archbold Biol. Sta., 1-7.v.1961, R. W. Hodges, light trap, 1 male, 1 female;
13-19.iv.1970, W. W. Wirth, light trap, 5 males, 3 females. Sebring, Highlands Hammock
State Park, 15.iv.1970, W. W. Wirth, light trap, 1 male, 10 females. Hillsborough Co.,
Harris Swamp, 13.iv.1967, ? collector, 1 male. Indian River Co., Vero Beach, iv.1960,
Env. Res. Ctr. light trap, 1 female. Liberty Co., Torreya State Park, 15.iv.1957,
498
December, 1991
Wirth: Western Hemisphere Allohelea 499
27.iv.1958, F. S. Blanton, light trap, 8 males, 41 females; 20.v.1966, H. V. Weems, Jr.,
light trap, 3 males, 25 females; 22.iv.1967, W. W. Wirth, light trap, 3 males, 3 females.
Manatee Co., Cortez, ix.1960, St. Bd. Health, light trap, 1 male. Marion Co., Juniper
Springs State Park, 28.iv.1970, W. W. Wirth, light trap, 4 males, 6 females. Putnam
Co., Welaka, 9.iv.1964, H. A. Denmark, 2 females. Santa Rosa Co., Blackwater River
Biol. Sta., 21.v.1971, G. B. Fairchild, UV light trap, 1 female; Jay, v.1962, T. W. Boyd,
light trap, 1 male. Wakulla Co., Ochlockonee River State Park, 29.iv.1970, W. W. Wirth,
light trap, 4 males, 6 females. KANSAS: Douglas Co., Lawrence, v.1956, A. R. Barr,
light trap, 1 female. MARYLAND: Baltimore Co., Green Spring Valley, 14.vi.1952, R.
H. Foote, 1 female. Garrett Co., Swallow Falls State Park, 21.vii.1983, P. G. Bystrak,
light trap, 1 female. Montgomery Co., Colesville, 17.vi-18.vii.1975, W. W. Wirth, light
trap, 2 females; Fairland, 10,23.vi.1959, A. A. Hubert, light trap, 1 male. Prince Georges
Co., Patuxent Wildlife Res. Ctr., 16.vii.1979, W. W. Wirth, malaise trap, 1 female;
vi.1976, vii.1978, vii.1979, W. L. Grogan, Jr., malaise trap, 10 females. Wicomico Co.,
Salisbury, vi-viii.1978, vi.1981, W. L. Grogan, Jr., malaise trap, 1 male, 2 females.
MICHIGAN: Cheboygan Co., 17.vii.1942, C. W. Sabrosky, 1 female (paratype); Douglas
Lake, vii.1954, R. W. Williams, light trap, 4 males, 1 female. NORTH CAROLINA:
Jackson Co., Dulaney Bog, 10 km S Cashiers, vii.1987, W. W. Wirth, malaise trap, 1
female. Macon Co., Highlands, 22.vi.1962, R. C. & A. Graves, reared from Boletus
fungus, 1 female; vi.1986, vii.1987, W. W. Wirth, malaise trap, 4 males, 1 female.
Transylvania Co., Lake Toxaway, vii.1989, W. W. Wirth, UV light trap, 2 males, 1
female. SOUTH CAROLINA: Georgetown Co., Hobcaw House, vi.1972, L. Henry,
light trap, 1 male, 2 females; Pawley's Island, 20.vi.1971, L. Henry, light trap, 1 female.
VIRGINIA: Alexandria, 15-24.vi.1951, W. W. Wirth, stream margin, 2 females;
4.vii.1951, W. W. Wirth, Osmunda bog, 1 female. Augusta Co., Mount Solon, 22.vi.1951,
W. W. Wirth, light trap, 2 females. Fairfax Co., Falls Church, 6.vii.1958, W. W. Wirth,
light trap, 1 male; Holmes Run, 3.vii.1960, W. W. Wirth, light trap, 1 female. Rockbridge
Co., Vesuvius, 13.vii.1960, D. H. Messersmith, 1 male.
Discussion. Allohelea johannseni is difficult to separate from A. nebulosa; distin-
guishing characters are listed above under nebulosa.
Allohelea distortifemur Wirth, new species
(Figs. 8-10, 16, 25)
Closely resembling Allohelea nebulosa (Coquillett), differing as described below;
female readily distinguished by the greatly constricted proximal half of the hind femur
with a distinct ventral knob-like protuberance at proximal third.
Female Holotype. Wing length 1.45 mm; breadth 0.53 mm; costal ratio 0.78.
Head: Brownish, proximal antennal segments and third palpal segment paler. An-
tenna (Fig. 8) with lengths of flagellar segments 65-36-43-43-43-43-47-47-72-72-72-72-86;
antennal ratio 0.97. Palpus (Fig. 9) with lengths of segments 25-32-72-54-75; palpal ratio
2.0.
Thorax: Uniformly dark brown, pattern of pollinosity not visible in slide preparations.
Legs without yellowish knee spots; all tarsi stramineous; fore and mid femora and tibiae
pale brown, slender. Hind femur and tibia blackish, greatly thickened; femur (Fig. 10)
with a sharp constriction just before midlength, and proximad of this a prominent bulbous
ventral swelling; lengths of segments from femur to T5, 900-860-300-165-108-122-108;
hind claw 144, basal tooth 43. Wing with pattern of dark areas as in nebulosa. Halter pale.
Abdomen: Dark brown, proximal segments somewhat yellowish; cerci yellowish.
Spermathecae (Fig. 20) large and globular, irregularly subspherical to short oval, with
long, gradually tapering neck; slightly unequal, 144 x 100 and 118 x 93 with necks 22;
vestigial spermatheca threadlike.
Florida Entomologist 74(4)
Figs. 20-23. Allohelea spp., male genitalia: 22, bottimeri; 23, nebulosa; 24, neotropica;
25, distortifemur; 26, weemsi.
Male Allotype. Wing length 144 mm; breadth 0.38 mm; costal ratio 0.75.
Similar to female, with usual sexual differences; hind femur not modified, only slightly
narrowed toward base. Hind leg with lengths from femur to T5, 900-860-289-172-72-100-
100; claw 115, basal tooth 22.
Genitalia (Fig. 25): Similar to those ofA. nebulosa; distinguishing features: Aedeagus
with proximal sclerite slightly broader than long; distal sclerite short and stout, the
lateral sheaths deeply pigmented and forming a bulbous lateral swelling proximally.
Dististyle short, 0.50 as long as basistyle, curved, and tapering to sharp distal point.
Parameres with distal processes only slightly curved, sinuate, and not curved dorsally
around the conically tapering ninth tergum.
500
December, 1991
Wirth: Western Hemisphere Allohelea
Distribution. Florida, known presently only from the type locality.
Types. Holotype female, allotype male, 14 male and 20 female paratypes, Florida,
Gulf Co., St. Joseph State Park, 1-3.v.1970, W. W. Wirth, UV light trap.
Discussion. The specific epithet is from the Latin: distortus,-misshapen or deformed,
referring to the greatly modified hind femur of the female. No similar modification is
known in any other species of Allohelea. The male is readily distinguished from other
American Allohelea by the short, pointed dististyle.
Allohelea weemsi Wirth, new species
(Figs. 11, 26)
Closely resembling Allohelea nebulosa (Coquillett), differing as described below;
female readily distinguished by its large size (wing length 1.65 mm), and greatly swollen
hind femur, swollen nearly to the base.
Female Holotype. Wing length 1.65 mm; breadth 0.60 mm; costal ratio 0.78.
Head: Brownish, bases of antennal segments 3-10 and all of third palpal segment
yellowish. Antenna with lengths of flagellar segments 57-54-54-54-54-54-54-54-86-86-93-
97-129; antennal ratio 1.13, distal five segments greatly elongated. Palpus with lengths
of segments 25-40-79-58-72; palpal ratio 2.7.
Thorax: Dark brown, pollinose pattern not visible in slide preparations, darker brown
dots at seta bases; scutellum paler in midportion. Fore and mid legs dark brown, slender;
tarsi stramineous. Hind leg with femur and tibia greatly swollen, blackish; tarsus pale
brown, the joints narrowly blackish; hind femur (Fig. 11) with a short basal constriction,
greatly and evenly swollen to apex, breadth 180; hind tibia also swollen but less so,
breadth 152; lengths of segments from femur to T5, 935-935-340-187-108-130-115; hind
claw 160, basal tooth 51. Wing with pattern of dark areas as in A. nebulosa. Halter pale.
Abdomen: Dark brown, proximal segments paler to yellowish; cerci pale brownish.
Spermathecae as in A. pedicellata, large and strongly sclerotized; measuring 122 x 94
and 115 x 87 not including necks; necks slender, not tapering, 10 microns long; vestigial
spermatheca short and slender.
Male Allotype. Wing length 1.70 mm; breadth 0.52 mm; costal ratio 0.81.
Similar to female with usual sexual differences. Antenna with segments 3-10 and
plume pale. Dark area of wing paler and less extensive. Hind femur less swollen, gradu-
ally narrowing to base; lengths from femur to T5, 1040-1001-360-241-94-144-136; claw
165, basal tooth 36.
Genitalia (Fig. 26): Similar to those of A. nebulosa; distinguishing features: Dististyle
short, curved, and markedly tapering to bluntly pointed tip; only 0.58 as long as basistyle.
Aedeagus with slender, slightly tapering distal peglike internal sclerotization, flanked
by a more lightly sclerotized structure with parallel sides at base, distally expanded in
a large arrowhead-shaped tip that is deeply cleft proximally on dorsal side as figured.
Parameres with bulbous basal portion short, heavily sclerotized distally, tapering and
markedly curving to hornlike process curving dorsomesad around ninth tergum.
Distribution. Florida.
Types. Holotype female, allotype male, Highlands Hammock State Park, Highlands
Co., Florida, 15.iv.1970, W. W. Wirth, UV light trap. Paratypes, 3 males, 8 females
as follows: FLORIDA: Alachua Co., Gainesville, Chantilly Acres, 7-8.v.1967, F. S.
Blanton, light trap, 2 females. Highlands Co., same data as type, 2 females. Liberty
Co., Torreya State Park, 20.v.1966, H. V. Weems, Jr., 1 female. Santa Rosa Co.,
Blackwater River Biol. Sta., 21.v.1971, G. B. Fairchild, UV light trap, 3 males, 3 females.
Discussion. This species is dedicated to Howard V. Weems, Jr., in recognition of his
outstanding contributions to the taxonomy of Florida Diptera as the prodigious collector
and for 38 years the indefatiguable curator of the Florida State Collection of Arthropods.
Florida Entomologist 74(4)
Allohelea weemsi can be distinguished from its Florida congeners by its greatly
swollen female hind femur, abruptly narrowed only at its base, by the high antenna
ratio (1.13), by the enlarged, oval spermathecae with slender necks, by the short tapering
male dististyle, and by the sagittate shape of the distal portion of the male aedeagus.
Next to A. pedicellata it is the largest of the North American species of Allohelea.
Allohelea pedicellata Wirth, new species
(Figs. 12, 21)
Closely resembling Allohelea nebulosa (Coquillett), differing as described below;
female readily distinguished by the greatly and abruptly constricted, subcylindrical,
proximal third of the hind femur.
Female Holotype. Wing length 1.69 mm; breadth 0.59 mm; costal ratio 0.80.
Head: Brown, third palpal segment paler. Antenna with lengths of flagellar segments
54-40-43-43-47-47-47-50-80-83-83-83-90; antennal ratio 1.12. Palpus with lengths of seg-
ments 18-40-72-43-58; palpal ratio 2.5.
Thorax: Scutum pruinose gray with black dots at seta bases; scutellum dark brown.
Legs without yellowish knee spots; fore and mid tarsi stramineous. Hind femur and
tibia blackish, hind tarsus brown; hind femur with abrupt, subcylindrical constriction
on proximal third (Fig. 12); lengths of segments from femur to T5, 970-915-340-165-108-
144-122; hind claw 165, basal tooth 57. Wing with pattern of dark areas as in nebulosa.
Halter pale.
Abdomen: Dark brown, proximal segments paler; cerci yellowish. Spermathecae
(Fig. 21) oval with short, slender necks; subequal, each measuring 125 x 86, with neck 18.
Male. Unknown.
Distribution. Western Florida, southern Alabama, and Georgia. Types. Holotype
female, Florida, Baker Co., Olustee, vii.1971, F. S. Blanton, UV light trap. Paratypes,
13 females. ALABAMA: Baldwin Co., Gulf Shores, Gulf State Park, 24-27.v.1978, J. I.
Glick, 1 female. FLORIDA: Baker Co., same data as type, 1 female. Jackson Co.,
Florida Caverns State Park, 26.v.1973, W. W. Wirth, UV light trap, 1 female. Jefferson
Co., Monticello, iv.1969, W. H. Whitcomb, UV light trap, 1 female. Leon Co., Tall
Timbers Res. Sta., 29.v.1973, W. W. Wirth, UV light trap, 2 females. Liberty Co.,
Torreya State Park, 27.iv.1958, F. S. Blanton, light trap, 3 females; 20.v.1966, H. V.
Weems, Jr., 2 females. Santa Rosa Co., Jay, v.1962, T. W. Boyd, light trap, 1 female.
GEORGIA: Thomas Co., Thomasville, 15-30.v.1948, E. Palmer, light trap, 1 female.
Discussion. This is the largest of the North American species of Allohelea. It is
readily distinguished in the female by the subcylindrical basal constriction of the hind
femur, from which the species takes its name: pedicellus (Latin),- small slender stalk.
I have not been able to identify the male of this species in available material; it may be
inseparable from the male of A. nebulosa.
Allohelea bottimeri Wirth, new species
(Figs. 18, 22)
Monohelea nebulosa (Coquillett), in part; Wirth & Williams, 1964:304 (misident.; records
from Kerrville, Texas).
A small species of Allohelea similar to the Palaearctic A. tesselata (Zetterstedt), but
the genitalia with short dististyle with blunt tip and the parameres with much shorter
caudal processes, the tips of which are not recurved.
Female Holotype. Wing length 1.19 mm; breadth 0.45 mm; costal ratio 0.75.
December, 1991
Wirth: Western Hemisphere Allohelea 503
Head: Brown including antennae and palpi. Antenna with lengths of flagellar seg-
ments 58-40-36-40-41-41-41-41-62-62-62-62-79; antennal ratio 0.89. Palpus with lengths
of segments 18-47-72-36-61; palpal ratio 2.9.
Thorax. Dark brown; scutum without prominent pattern. Wing as in A. johannseni;
halter pale. Legs without yellowish knee spots, femora and tibiae brown, blackish on
hind leg; tarsi stramineous. Hind femur moderately swollen, slightly and gradually
tapering to base, as in A. johannseni (Fig. 14). Hind leg with lengths of segments from
femur to T5 720-680-260-144-72-108-86; claw 125, basal tooth 43.
Abdomen: Dark brown to base, cerci pale. Spermathecae (Fig. 18) short oval with
distinct slender necks; unequal, 69 x 52 and 58 x 48, with necks 14 microns long.
Male Allotype. Wing length 1.11 mm; breadth 0.35 mm; costal ratio 0.78.
A uniformly dark brown species as seen in slide preparations; hind femur and tibia
blackish, yellow knee spots absent; tarsi stramineous. Antenna with lengths of segments
12-15, 54-111-101-93. Hind leg with lengths from femur to T5, 723-680-237-136-65-93;
claw 122, basal tooth 22.
Genitalia (Fig. 22): Similar to those of A. nebulosa, but ninth tergum short and
broad, distal portion tapering to widely spaced, prominent, angular apicolateral proces-
ses. Basistyle about twice as long as greatest breadth; dististyle short, 0.65 as long as
basistyle, stout, nearly straight on distal portion, with stout, blunt tip. Aedeagus about
as broad as long; basal sclerite nearly twice as broad as long; distal sclerite a broad,
roughly triangular plate with slender terminal process and rounded lateral shoulders
bearing minute serrations; ventral surface with distinct longitudinal striations. Para-
meres short and relatively stout, mesal connection not visible in available preparations;
posterior portion bladelike with bluntly pointed apex abruptly bent dorsomesad.
Distribution. Texas.
Types. Holotype female, Kerrville, Kerr Co., Texas, 21.v.1954, L. J. Bottimer, light
trap. Allotype male, same data, but iv.1954. Paratypes, 5 males, 19 females, as follows:
TEXAS: Kerr Co., Kerrville, iv.1954, L. J. Bottimer, 4 males, 8 females; same data
but v.1954, 10 females; same but vi.1954, 1 female. Dallas Co., Dallas, 5.v.1942, H.
Knutson, 1 male.
Discussion. This species is dedicated to the late Larry D. Bottimer in appreciation
of his friendship and unselfish assistance during several years of field work on the biting
midges of West Texas.
Allohelea bottimeri appears to be most closely related to the Palaearctic species
Allohelea tesselata (Zetterstedt), the male of which was described by Wirth (1953). In
tesselata the male dististyle is much longer, curved and slender distally as in typical
Allohelea, while the parameres were figured with straight stems and longer recurved
tips. In North America, A. bottimeri closely resembles A. johannseni, but is distin-
guished by its more uniformly dark brown body, by the short, stout, male dististyle,
and by the short, broad, distal portion of the aedeagus.
Allohelea neotropica Wirth, new species
(Figs. 15, 17, 24)
Monohelea johannseni Wirth; Lane & Wirth, 1964: 214 (misident.; Panama records).
A small dark brown species resembling Allohelea johannseni (Wirth), differing as
described below; hind femur with apical yellow knee spot.
Female Holotype. Wing length 0.99 mm; breadth 0.54 mm; costal ratio 0.81.
Head: Brownish; palpal segments 2-4 pale. Antenna with lengths of flagellar segments
54-36-36-36-36-36-36-36-58-61-72-65-76; antennal ratio 0.89. Palpus with lengths of seg-
ments 18-25-47-32-57; palpal ratio 1.8.
504 Florida Entomologist 74(4) December, 1991
Thorax: Dark brown; scutum subshining brown with gray pollinose areas containing
blackish dots at seta bases; scutellum gray pollinose, narrowly yellowish on midportion.
Legs brown, hind femur and tibia blackish; knee spots yellowish, prominent on hind
leg; tarsi stramineous. Hind femur and tibia greatly thickened, femur (Fig. 15) uniformly
stout to base as in johannseni; lengths of segments from femur to T5, 610-540-193-93-65-
86-86; claw 144, basal tooth 43. Wing with pattern of dark areas as injohannseni. Halter
pale.
Abdomen: Dark brown, proximal segments paler; cerci yellowish. Spermathecae
(Fig. 17) small, ovoid with moderately long, tapering necks; unequal, measuring 62 x
46 and 58 x 38, with necks 15.
Male Allotype. Wing length 1.09 mm; breadth 0.35 mm; costal ratio 0.72.
Similar to female with usual sexual differences. Antenna with lengths of flagellar
segments 94-36-36-36-36-36-36-36-36-47-108-79-90. Hind leg with lengths from femur to
T5, 590-530-186-100-72-86-86; claw 115, basal tooth absent.
Genitalia (Fig. 24): Slightly longer than broad, not so broad as in the Nearctic species,
closely resembling those of A. johannseni but dististyle longer and distally more slender;
aedeagus with basal sclerite about twice as broad as long and distal sclerite with distinct
apical notch.
Distribution. Neotropical; known from Jamaica, Belize, Panama, and Colombia.
Types. Holotype female, allotype male, Jamaica, Westmoreland Parish, Negril, Crys-
tal Waters, 20.xi.1968, R. E. Woodruff, UV light trap in tropical hammock. Paratypes,
6 males, 9 females: JAMAICA: Same data as holotype, 3 males. Negril Beach, 12.iii.1970,
W. W. Wirth, rocky shore, 1 male, 2 females. Trelawny Parish, 2 mi N Troy, 26.viii.1969,
R. E. Woodruff, 1 female. Hardwar Gap, Hollywell, 16.vi.1970, E. G. Farnworth, light
trap, 2 females; Green Hills, Inst. Jamaica cabin, 18.viii.1969, R. E. Woodruff, 2 males,
2 females. BELIZE: 1.5 mi W Punta Gorda, 31.vii.1968, W. L. Haase, UV light trap,
1 male. COLOMBIA: Valle, Rio Raposo, viii.1964, V. H. Lee, light trap, 1 female.
PANAMA: Bocas del Toro Prov., Almirante, 28.x.1952, i.1953, F. S. Blanton, light
trap, 1 male, 1 female. Canal Zone, Albrook Field, 19.vi.1952, F. S. Blanton, light trap,
1 female; Barro Colorado Id., vii.1967, W. W. Wirth, light trap, 1 female; Mojinga
Swamp, Fort Sherman, 25.vi.1952, F. S. Blanton, 1 male.. Los Santos Prov., Panama
de Azucar, 10.x.1952, F. S. Blanton, light trap, 1 female.
Discussion. Allohelea neotropica was misidentified by Lane & Wirth (1964) from
Panama as Monohelea johannseni. It is the smallest of the American species and is also
distinguished by the prominent yellow knee spots. The male dististyle is longer and
distally more slender than in johannseni, and the proximal sclerite of the aedeagus is
short and broad.
ACKNOWLEDGMENTS
I am grateful to Molly K. Ryan for the habitus drawing of Allohelea nebulosa (Coquil-
lett) (Fig. 1). I also am grateful to William L. Grogan, Jr., Salisbury State University,
Salisbury Maryland, Art Borkent, Salmon Arm, British Columbia, and J. Antony
Downes, Ottawa, Ontario, Canada, for careful and constructive review of an earlier
draft of the manuscript. Acknowledgment is also made to the following persons and
institutions for loan of specimens: Auburn University, Auburn, Alabama, courtesy of
Gary Mullen and Steven Murphree; and U.S. National Museum of Natural History,
courtesy of Ronald McGinley and B. V. Peterson (Systematic Entomology Laboratory,
USDA).
Wirth: Western Hemisphere Allohelea
REFERENCES CITED
COQUILLETT, D. W. 1901. New Diptera in the U. S. National Museum. Proc. United
States Natl. Mus. 23: 593-618.
DOWNES, J. A., AND W. W. WIRTH. 1981. Ceratopogonidae, pp. 393-421, in McAlpine,
J. F., et al. [eds.], Manual of Nearctic Diptera. Vol. 1. Agriculture Canada
Monograph No. 27, 674 pp.
GRAVES, R. C., AND C. F. GRAVES. 1985. Diptera associated with shelf fungi and
certain other micro-habitats in the Highlands area of western North Carolina.
Entomol. News 96: 87-92.
JOHANNSEN, 0. A. 1943. A generic synopsis of the Ceratopogonidae (Heleidae) of
the Americas, a bibliography, and a list of the North American species. Ann.
Entomol. Soc. America 36: 763-791.
KIEFFER, J. J. 1906. Diptera. Fam. Chironomidae. Fasc. 42, 78 pp., 4 plates, in
Wystman, P. [ed.], Genera Insectorum. Bruxelles.
KIEFFER, J. J. 1917. Chironomides d'Amerique conserves au Musee National Hongrois
de Budapest. Budapest Magyar Nemzeti Muz., Ann. Hist. Nat. 15: 292-364.
LANE, J., AND W. W. WIRTH. 1964. The biting midge genus Monohelea Kieffer in
the Neotropical Region (Dipt. Ceratopogonidae). Studia Entomol. 7: 209-236.
MALLOCH, J. R. 1914. Notes on North American Diptera, with descriptions of new
species in the collection of the Illinois State Laboratory of Natural History. Bull.
Illinois St. Lab. Nat. Hist. 10: 213-243, 3 plates.
MALLOCH, J. R. 1915. The Chironomidae, or midges, of Illinois, with particular refer-
ence to the species occurring in the Illinois River. Bull. Illinois St. Lab. Nat.
Hist. 10: 275-543, 23 plates.
WIRTH, W. W. 1953. American biting midges of the heleid genus Monohelea. Proc.
United States Natl. Mus. 103: 135-154.
WIRTH, W. W., AND W. L. GROGAN, JR. 1981. Natural History of Plummers Island,
Maryland XXV. Biting midges (Diptera: Ceratopogonidae). 3. The species of the
tribe Stilobezziini. Bull. Biol. Soc. Washington no. 5: 1-102.
WIRTH, W. W., AND W. L. GROGAN, JR. 1988. The predaceous midges of the world
(Diptera: Ceratopogonidae; Tribe Ceratopogonini). Flora & Fauna Handbook no.
4, 160 pp. E. J. Brill, Leiden.
WIRTH, W. W., AND R. W. WILLIAMS. 1964. New species and records of North
American Monohelea (Diptera: Ceratopogonidae). Ann. Entomol. Soc. America
57: 302-310.
505
Florida Entomologist 74(4)
FORCIPOMYIA BICOLOR AND RELATED SPECIES OF
THE SUBGENUS LEPIDOHELEA IN BRAZIL
(DIPTERA: CERATOPOGONIDAE)
WILLIS W. WIRTH
Research Associate, Florida State Collection of Arthropods
1304 NW 94th St., Gainesville, Florida 32606, U.S.A.
ABSTRACT
Forcipomyia (Lepidohelea) bicolor Lutz from Brazil and two other closely related
species, F. dubia Macfie and F. lacrimotorii Macfie, are redescribed from types. For-
cipomyia discoloripes Macfie is a junior synonym of F. bicolor (NEW SYNONYMY).
The Forcipomyia bicolor Group of species is diagnosed and a revised key is presented
to the nine described Neotropical species.
RESUME
En base a prototipos se describe de nuevo la especie Foripomya (Lepidohelea) bicolor
Lutz del Brazil y otras 2 species estrechamente relacionadas, F. dubia Macfie y F.
lacrimotorii Macfie. Forcipomyia discoloripes Macfie es un sinonimo de F. bicolor
(Nuevo Sinonimo). Se hace una diagnosis del grupo de species de Forcipomyia bicolor
y se present una clave revisada describiendo las nueve species neotropicales.
After nearly 50 years of investigation, the most important pollinators of Theobroma
cacao L., source of commercial chocolate and cocoa, have been found to belong to the
ceratopogonid genus Forcipomyia Meigen (Billes 1941, Macfie 1944, Posnette 1950,
Saunders 1956, 1959, Winder 1978, Young 1983). Earlier research indicated that the
most frequent and important pollinators were species of the subgenus Euprojoannisia
Brethes (Saunders 1959, Soria & Wirth 1974, Bystrak & Wirth 1976). More recently
attention has also turned to members of other subgenera of Forcipomyia as more abun-
dant in cacao plantings and also important as cacao pollinators. Thus Winder (1977,
1978) found that about a third of the Forcipomyia midges in cacao groves in Bahia,
Brazil, belonged to the subgenus Forcipomyia s. str.; Young (1983) found the proportion
much higher in Costa Rica. Kaufmann (1975), however, found F. (Microhelea) inor-
natipennis (Austen) and F. (Lepidohelea) squamipennis Ingram & Macfie to be more
important in Ghana.
Winder (1978) pointed out that there are so many variables in weather conditions,
available terrestrial midge habitats, and flowering patterns of the cacao trees themselves,
that relative importance of midges and other insects as pollinators is difficult to evaluate.
Moreover, cage experiments with insects on cacao flowers also usually lead to unrealistic
results. Nevertheless, recent collections that I have examined made by John Winder in
the cacao plantation Fazenda Almirante in Bahia, Brazil, show that several species of
the subgenus Lepidohelea Kieffer are abundant and at times dominant in cacao planta-
tions.
The systematics of the subgenus Lepidohelea remained extremely confused until
Debenham (1987), in reviewing the Australasian species of the genus Forcipomyia,
succeeded in finding reliable adult characters to separate species of Lepidohelea from
the closely related Forcipomyia s. str. Species of Lepidohelea can be recognized by
their spindle-shaped third palpal segment, legs with characteristic banding of alternating
December, 1991
Wirth: Forcipomyia bicolor Group 507
pale and dark bands, at least on hind tibia, male dististyle without long setae on the
outer margin, and the larva with b hairs of body and p and q hairs swollen near base
and becoming filamentous distally. Wirth (1991) followed with a key to divide the species
of Lepidohelea into three species groups as follows:
1. Palpus with four segments; one spermatheca; male dististyle more or less
expanded distally (Western Hemisphere species) ............. annulatipes Group
1'. Palpus with five segments; two spermathecae; male dististyle various ...... 2
2(1). Male dististyle straight or slightly curved, tapering to slender tip (Western
Hemisphere species) ........................................................ bicolor Group
2'. Male dististyle straight to sinuate, tip more or less expanded (Eastern
Hemisphere species) ................................................. chrysolopha Group
The species of the chrysolopha Group were admirably discussed and the Australasian
species revised by Debenham (1987). The annulatipes Group has not been formally
diagnosed, but is now known to comprise three Brazilian species, annulatipes Macfie,
brasiliensis Macfie, and kuanosceles Macfie (Macfie 1939). I now have in preparation a
revision of this group which is also represented in North America by at least two
undescribed species. Wirth (1991) did not diagnose the bicolor Group, but presented a
key to nine Neotropical species, of which one, F. winderi, was described as new. The
bicolor Group is represented in North America by four described species, beckae Wirth,
christiansoni Wirth & Hubert, seminole Wirth, and varipennis Wirth & Williams, and
at least six undescribed species. The immature stages of members of the bicolor Group
have not been described.
Because of their abundance in collections of cacao pollinators, it is urgent that accurate
names be ascribed to the important pollinators and that keys be presented for their
identification. As a preliminary to planned revisions, first of the Nearctic species, and
then of the Neotropical species, of the bicolor Group, it was necessary to study the
available types of previously described species, whose identity remained questionable
because of inadequate descriptions and figures. Notes are presented here on four Brazi-
lian species described by Lutz (1914) and Macfie (1939): bicolor Lutz, discoloripes Macfie,
dubia Macfie, and lacrimotorii Macfie. The remaining six previously described Neotrop-
ical species, abercrombyi Macfie, flavifemoris Macfie, seminole Wirth, squamithorax
Clastrier, varipennis Wirth & Williams, and winderi Wirth, will be treated elsewhere
when adequate taxonomic material is assembled. As a result of the present study a
diagnosis is presented for the bicolor Group, a new synonymy is discovered, three
species are redescribed and figured, and a corrected key is devised for the identification
of the described Neotropical species.
Explanation of the taxonomic characters used can be found in the general papers on
Ceratopogonidae by Wirth et al. (1977) and Downes & Wirth (1981), and the revision
of the North American Euprojoannisia by Bystrak & Wirth (1978).
Forcipomyia bicolor Group
Diagnosis.-Medium size to large species of Forcipomyia with legs more or less pale
banded (Fig. 1). Wing length 0.7-1.5 mm. Female wing often with mottled pattern of
pale areas; male wing usually extensively pale with restricted dark areas. Body with
flattened scales, from long and 1-striated to broad and flattened and multi-striated.
Head, body and legs with abundant semi-erect setae, these often long and bristly.
Female antenna short to moderately long, segments vase-shaped to tapering; no sharp
division in length between segments 10 and 11. Male antenna with plume usually yellowish
Florida Entomologist 74(4)
December, 1991
.-ie- .
-:Y^^^^^i:i; Wfi .1
'+,........ .............. ::....... ':'.
+ -5. . .
S::~ ^ MJ
------- --- ___ ______ '* **'* '
BICOLOR
DUBIA
LACRIMOTORII
ABERCROMBYI
WINDERI
FLAVIFEMORIS
SEMINOLE
VARIPENNIS
Fig. 1. Diagram of pale and dark markings of femora (left) and tibiae) right) of (top
to bottom) fore, mid, and hind legs of species of the Forcipomyia bicolor Group.
apically. Palpus 5-segmented; third segment variably swollen in midportion, sensory pit
small, shallow to deep. Female mandible without teeth. Hind tibia always with pale
bands; femora and other tibiae with variable markings to unbanded. Hind tarsal ratio
about 1.0. Tarsal claws curved and slender; outer claw of male fore tarsus often with
508
Wirth: Forcipomyia bicolor Group 509
blunt ventral tooth at midlength (Fig. 9). Female with two spermathecae. Female genital
sclerotization (Fig. 7) usually an arcuate transverse ribbon bearing small spines on
posterior margin, especially at lateral ends. Male genitalia (Fig. 11) often bicolored;
dististyle not swollen or modified at tip. Aedeagus (Fig. 10) usually with low basal arch,
more or less triangular in outline, and bearing 1-3 inconspicuous longitudinal ridges on
ventral surface. Parameres not joined at base, consisting of simple, long, nearly straight
rods tapering to filiform tip.
Revised Key to Neotropical Species of the
Forcipomyia bicolor Group
(Fig. 1)
1. Femora dark from base to tip (narrow knee spots may be pale, narrow
bases may be pale) ............................................. ..................... 2
1' Femora extensively pale proximally, may be entirely pale ....................... 5
2(1). All tibiae entirely dark; (antennae long, segments with narrow base swol-
len, abruptly narrowed to long slender distal portion; mesonotum with
four pairs of lines of white scales) ......................... squmithorax Clastrier
2'. At least hind tibia with broad sub-basal pale band .................................. 3
3(2). All tibiae with pale band on proximal half ........................................... 4
3'. Fore and mid tibiae entirely dark; hind tibia with sub-basal and apical pale
bands ............................................................. dubia M acfie
4(3). Mesonotum with sublateral pair of compact tufts of long black setae; male
aedeagus twice as long as basal breadth, basal arch to 1/5 of total length ...........
.............................................. .............. lacrim otorii Macfie
4'. Mesonotum without discal tufts of long black setae; male aedeagus short with
broad base, total length subequal to basal breadth, basal arch to nearly half
of total length ........................................................ ... abercrombyi Macfie
5(1). Femora pale except median brown band on hind femur ........... seminole Wirth
5'. Femora with distal brown bands, at least on hind leg ................................ 6
6(5). Femora dark on distal half or more; third palpal segment of female greatly
swollen to past midportion, with deep pit; wing markings, tibial spur, and
m ale fore claw various ............................................................................. 7
6'. Femora pale or with narrow basal and/or apical brown bands on mid and/or
hind legs; third palpal segment of female moderately swollen with shallow
pit; female wing with pale markings; spur of hind tibia short and nearly
straight; male fore tarsus without ventral tooth on one claw ....................... 8
7(6). Female wing uniformly dark brown; spur of hind tibia long, curved; one claw
of male fore tarsus with blunt ventral tooth at midlength .............. bicolor Lutz
7'. Female wing mottled with pale markings; spur of hind tibia short and
near straight; male fore tarsus without ventral tooth on one claw .............
......................................... ............................t varipennis W irth & W illiams
8(6). Fore and mid femora entirely pale; hind femur pale with broad apical
brown band .................................................... winderi W irth
8'. Fore and mid femora with narrow basal brown band; hind femur with
narrow apical brown band ............................................. flavifemoris Macfie
Forcipomyia (Lepidohelea) bicolor Lutz
(Figs. 2-11)
Forcipomyia bicolor Lutz, 1914: 89 (male; Brazil).
Forcipomyia (Forcipomyia) bicolor Lutz; Wirth, 1974: 5 (catalog).
510 Florida Entomologist 74(4) December, 1991
72 3
36 1` 6
72
7 125
9
10
Figs. 2-11. Forcipomyia bicolor; 2, 5-7, female from Santa Catarina, Brazil
(Plaumann); 3-4, 8-11, lectotype male, Santa Catarina, Brazil (Plaumann): 2, antenna
segments 7-15; 3, antennal segments 12-15; 4, 6, palpus; 5, spermathecae; 7, genital
sclerotization; 8, apex of tibia and basitarsus of hind leg; 9, fifth tarsomere and claws
of fore leg; 10, aedeagus; 11, genitalia, aedeagus and one dististyle omitted (scale in
microns).
Forcipomyia discoloripes Macfie, 1939: 159 (male, female; Brazil; fig. female antenna,
palpus). NEW SYNONYMY
Forcipomyia (Forcipomyia) discoloripes Macfie; Wirth, 1974: 5 (catalog).
A large species (wing length 1.30 mm in female) with uniformly dark brown wings in
female, wing grayish with two distinct dark spots in male; femora dark brown on distal
two-thirds; female palpus greatly swollen with deep sensory pit; hind tibial spur long and
curved; outer claw on male fore tarsus with blunt ventral tooth at midlength.
Female Plesiotype.-Wing length 1.39 mm; breadth 0.52 mm; costal ratio 0.47.
Head: Dark brown including antenna and palpus; long blackish bristles arching over
eyes. Antenna (Fig. 2) with lengths of flagellar segments 72-58-58-61-61-61-61-61-82-79-79-
79-93 microns; antenna ratio 0.85; segments 4-10 short and somewhat flask-shaped, 11-15
longer and tapering distally; whorls of sensilla chaetica on 3-10 strong and bristle-like.
Wirth: Forcipomyia bicolor Group 511
Palpus (Fig. 6) with lengths of segments 36-36-93-36-43 microns; third segment greatly
swollen on proximal 0.7, with a deep sensory pit opening by a smaller pore; palpal ratio 2.00.
Thorax: Dark brown; mesonotum with sparse, erect, dark brown bristles and sparse,
pale brown, short appressed hairs. Legs with dark brown and yellowish brown bands
as in Fig. 1; with rather long bristles, especially on tibiae, and numerous broad, ap-
pressed, striated scales. Spur of hind tibia long, curved and pubescent at base, as long
as distal breadth of tibia, pale on basal half, dark distally. Tarsi dark brown with narrow
pale rings at the joints, fifth tarsomeres pale; claws slender, curved, simple; hind tarsal
ratio 1.00. Wing uniformly dark brown, with dense decumbent dark brown macrotrichia.
Halter brownish.
Abdomen: Dark brown, with dense vestiture of decumbent, dark brown, single-
striated scalelike hairs, mixed on proximal segments with broad appressed scales; last
segment with long golden hairs. Genital sclerotization (Fig. 7) a narrow bowed ribbon
with ends slightly broadened and bearing 2-3 slender spines, the concave posterior
margin with three pairs of widely spaced, shorter, pale spines. Spermathecae (Fig. 5)
ovoid with slender necks; unequal, overall measuring 86 by 53 microns and 61 by 48
microns.
Male Lectotype.-Wing length 1.44 mm; breadth 0.36 mm; costal ratio 0.43.
As in female with usual sexual differences. Wing with pale gray macrotrichia except
two areas of darker, brownish macrotrichia, one over radial cells and the other over
vein Cul. Halter pale. Legs with more extensive pale markings and fainter dark markings
on fore and mid legs, their femora darker on about distal third and the dark bands on
tibiae narrower.
Antennal segments 12-15 (Fig. 3) with lengths 180-155-118-94 microns; plume brow-
nish proximally, yellowish distally. Palpus as in Fig. 4; palpal ratio 2.30. Fore tarsus
with outer claw (Fig. 9) broadened in midportion in a blunt tooth. Hind tibial spur (Fig.
10) greatly enlarged (length 115 microns) and curved, sickle-shaped, yellowish proximally
and gradually darkened toward tip. Hind basitarsus somewhat bowed and bearing strong
ventral spines; hind tarsal ratio 0.94.
Genitalia (Fig. 11): Brown, base of basistyle and midportion of ninth sternum yel-
lowish. Dististyle nearly straight. Aedeagus (Fig. 10) 1.4 times as long as basal breadth,
appearing almost triangular with basal arms short and basal arch not developed, anterior
margin well sclerotized, sides slightly convex; median ridge well-developed, lateral ridges
displaced to lateral margins. Parameres of usual form in bicolor Group; moderately
separated, the hyaline distal portions slender, slightly sinuate, and rather elongate,
their tips greatly surpassing tip of aedeagus.
Distribution.-Brazil, Costa Rica, Ecuador.
Types.-Forcipomyia bicolor Lutz (1914): "A descricao e feita de um macho montado
em preparado microscopic com gelatina glicerinada. Foi apanhado em Manguinhos num
aparelho de luz." Through the kindness of Maria Luiza Felippe-Bauer, Lutz' types were
borrowed from the Instituto Oswaldo Cruz in Rio de Janeiro, Brazil. These were mounted
on four slides, apparently the work of Dr. A. da Costa Lima who studied the Lutz
collection of Ceratopogonidae in 1937, as each slide bears the initials "C.L."
1. (left label): "Instituto Oswaldo Cruz / N. 3689 / Divisao 29 / Caixa 123." (right
label): "Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz / Na Luz, Man-
guinhos / Prs. coll. Dr. A. Lutz / C.L." Under a large square cover slip is mounted one
male cleared but with original coloration, intact but with genitalia dissected off and not
present.
2. (left label): "Instituto Oswaldo Cruz / N. 3721 / Divisao 1 / Caixa 125." (right label):
"Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz o / terminalia do exemp.
512 Florida Entomologist 74(4) December, 1991
/ da lam. 3689 / na Luz Manguinhos / Dn. coll. Dr. A. Lutz / C.L." Genitalia mounted
under large square coverslip.
3. (left label): "Instituto Oswaldo Cruz / N. 3691 / Divisao 1 / Caixa 124." (right label):
"Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz / Na luz, Manguinhos /
Da coll. A. Lutz / C.L. prep. x, 1937." Specimen apparently cleared in caustic and clorless;
intact except genitalia not present.
4. (left label): "Instituto Oswaldo Cruz / N. 3690 / Divisao 30 / Caixa 123." (right
label): "Instituto Oswaldo Cruz / N. 3288 / Forcipomyia / bicolor Lutz / N. luz, Manguinhos
/ Da coll. A. Lutz / Do expmpl. lam. 3691 / C.L. prep x,1937." Male genitalia preparation
cleared and much flattened.
Slides nos. 1 and 2 as above are hereby selected as the LECTOTYPE of Forcipomyia
bicolor Lutz.
Types of F. discoloripes: Macfie's (1939) description is a combined description of the
male and female sexes. No types were designated, but 16 male and 17 female specimens
examined were listed from Nova Teutonia, Brazil, with dates from 29.vii.1936 to
17. v.1937. These specimens may be considered syntypes. Through the courtesy of Bruce
C. Townsend the British Museum (Nat. Hist.) loaned me two of Macfie's slides which
were marked "Holotype male" and "Allotype female." The holotype male is hereby
selected as LECTOTYPE. It bears the locality data: "Brasilien / Nova Teutonia/ 270
11' B 52 23' L / Fritz Plaumann / 29,vii,1936." It has been dissected and mounted under
four coverslips and is in poor condition, but what remains agrees in all respects with
the male lectotype specimen of F. bicolor Lutz, under which name F. discoloripes Macfie
becomes a junior synonym (NEW SYNONYMY).
Other Specimens Examined.-BRAZIL: Santa Catarina, Nova Teutonia, 7 males,
12 females collected by F. Plaumann, with dates as follows: ix.1962, x.1962, viii.1963,
ix.1965, vii.1966, viii-ix.1970, ix.1970 (USNM). COSTA RICA: Puntarenas Prov.,
Sabalito, viii.1952, F. S. Blanton, light trap, 1 male. ECUADOR: Quevedo, Pichilingue,
INAP, vi. 1978, J. Mendoza, reared from rotting vegetation, 5 males, 8 females (USNM).
Forcipomyia (Lepidohelea) dubia Macfie
(Figs. 12-19)
Forcipomyia dubia Macfie, 1939: 162 (male, female; Brazil).
Forcipomyia (Forcipomyia) dubia Macfie; Wirth, 1974: 5 (catalog).
A large species (wing length 1.26 mm in female) with uniformly dark wings in female,
wing whitish with two faint dark spots in male; legs dark brown, only hind tibia with
pale bands; legs without broad scales; female palpus moderately swollen with deep
sensory pit; hind tibial spur not modified; outer claw of male fore tarsus with blunt
ventral tooth at midlength.
Similar to Forcipomyia bicolor, but differing as follows:
Female Plesiotype.-Wing length 1.26 mm; breadth 0.49 mm; costal ratio 0.43.
Head: Antennae (Fig. 12) with lengths of flagellar segments 72-65-61-61-61-61-61-61-
65-65-72-72-90 microns; antennal ratio 0.78. Palpus (Fig. 15) with lengths of segments
25-42-97-36-43 microns; third segment moderately swollen, with deep sensory pit; palpal
ratio 2.24.
Thorax: Legs (Fig. 1) dark brown, narrow knee spots yellowish; all femora and fore
and mid tibiae entirely dark brown, hind tibia with two prominent pale yellowish bands
as in bicolor. Hind tibial spur slender, length 75 microns. Legs with abundant long,
narrow, appressed one-striated scales, broad flat scales absent. Hind tarsal ratio 1.10.
Wing uniformly dark brown as in bicolor, with abundant long dark-brown macrotrichia.
Halter brownish.
Wirth: Forcipomyia bicolor Group
15
72_ J_ I L
17 1
Figs. 12-19. Forcipomyia dubia: 12-13, 15-16, female from Santa Catarina, Brazil
(Plaumann); 14, 17-19, lectotype male, Santa Catarina, Brazil (Plaumann): 12, antenna
segments 7-15; 13, spermathecae; 14, apex of hind tibia; 15, palpus; 16, genital scleroti-
zation; 17, fifth tarsomere and claws of fore leg; 18, aedeagus; 19, genitalia, aedeagus
and one dististyle omitted (scale in microns).
Abdomen: Spermatheca (Fig. 13) larger, apparently subequal (one tipped in orienta-
tion with neck hidden), slightly ovoid, 94 by 72 microns. Genital sclerotization (Fig. 16)
as in bicolor but lateral spines short and relatively stout.
Male Lectotype.-Wing length 1.61 mm; breadth 0.47 mm; costal ratio 0.42. Head
damaged (measurements from plesiotype: antennal segments 12-15 with lengths 234-165-
119-119 microns; palpal segments with lengths 36-43-119-47-40 microns, palpal ratio
4.10). Hind tarsal ratio 0.93; hind tibial spur (Fig. 14) 79 microns long. Fore tarsus with
blunt ventral tooth on outer claw as in bicolor (Fig. 17). Wing with pale macrotrichia
except two dark spots of black macrotrichia, one over radial cells and other on vein M1.
Halter pale.
Genitalia (Fig. 19): Brown, basal half of basistyle, median portion of ninth sternum,
and all except tip of dististyle yellowish. Dististyle slender and curved. Aedeagus (Fig.
18) nearly triangular in ventral view, 1.5 times as long as basal breadth, apex acutely
pointed; basal arch low but distinct, anterior margin well sclerotized; a median ridge
and two lateral ridges well marked. Parameres as in bicolor Group; bases widely sepa-
rated, distal portions hyaline, slender, reaching tip of aedeagus.
Distribution.-Brazil.
Types.-Syntypes, 2 males, Nova Teutonia, Brazil, 16 and 17.ix.1936 (BMNH). I
have studied the two syntypes loaned through the courtesy of Bruce Townsend and the
British Museum (Nat. Hist.). The specimen with date 16.ix.1936 had been mounted on
513
514 Florida Entomologist 74(4) December, 1991
a slide under two coverslips, and bears a blue circular label "Syntype" and a label with
red border printed "Type." This male is hereby designated LECTOTYPE of Forcipomyia
dubia Macfie. The second male, with date 17.ix.1936, is pinned and badly damaged,
consisting essentially of the thorax and one wing, with the genitalia mounted on a plastic
strip preparation on the pin below the specimen.
Other Specimens Examined.-BRAZIL: Santa Catarina, Nova Teutonia, ix.1962,
F. Plaumann (BMNH slide 1966-590), 1 female (Plesiotype). One male (Plesiotype), same
data but date viii-ix. 1970, 1 male (USNM).
Forcipomyia (Lepidohelea) lacrimotorii Macfie
(Figs. 20-20)
Forcipomyia lacrimotorii Macfie, 1939:161 (male, female; Brazil; fig. antenna, palpus).
Forcipomyia (Forcipomyia) lacrimotorii Macfie; Wirth, 1974; 5 (catalog).
A moderately large species (wing length 0.99 mm in female) with uniformly dark
brownish wings in both sexes; mesonotum in both sexes with a prominent pair of tufts
of long black setae; legs dark brown, tibiae with sub-basal pale bands and hind tibia
with apex broadly pale; legs with abundant broad striated scales; female palpus slightly
swollen with small shallow sensory pit; hind tibial spur not modified; outer claw of male
fore tarsus with blunt ventral tooth at midlength.
Very similar to Forcipomuyia dubia Macfie, differing as follows:
Female Plesiotype.-Wing length 0.99 mm; breadth 0.39 mm; costal ratio 0.42.
Head: Antenna (Fig. 20) with lengths of segments 57-61-65-65-65-65-65-65-61-65-65-
72-86 microns; antennal ratio 0.69; all segments with bulbous bases, then narrowing and
slightly tapering distally, less so on distal five segments. Palpus (Fig. 21) slender;
segments with lengths 29-36-79-24-29; third segment slightly swollen on proximal half,
palpal ratio 3.10; sensory pit small and shallow.
Thorax: Dark brown; mesonotum with sparse long erect black bristles and abundant
small appressed flattened whitish hairs; a pair of dense tufts of moderately long black
hairs sublaterally at midlength, about 10 hairs in each tuft. Legs (Fig. 1) dark brown,
including all femora; knees with narrow pale ring on tibiae; tibiae with moderately
narrow sub-basal pale ring and hind tibia with moderately broad apex pale; femora and
tibiae with abundant broad, striated, appressed scales. Hind tibial spur (Fig. 23) 65
microns long, straight and slender. Wing dark brown, slightly shaggy appearance due
to abundant long scalelike decumbent macrotrichia, especially prominent and dense over
radial veins. Halter brownish.
Abdomen: Spermathecae (Fig. 24) small, deeply pigmented; ovoid with scarcely
apparent necks; slightly unequal, measuring 61 by 44 microns and 52 by 40 microns.
Genital sclerotization a slender, bowed, transverse ribbon as in bicolor, but ends not
expanded and with no well-developed spines on posterior margin.
Male Plesiotype.-Wing length 1.36 mm; breadth 0.40 mm; costal ratio 0.42.
Head brown; antennal plume brown at base, yellowish distally. Antennal segments
12-15 with lengths 161-169-130-122 microns. Palpal segments with lengths 29-36-100-29-
30 microns; palpal ratio 1.60. Mesonotum with pair of prominent dark brown hair tufts
as in female. Hind tibial spur 90 microns long, straight and slender. Hind tarsal ratio
1.08. Fore tarsus with blunt ventral tooth on outer claw as in bicolor and dubia (Fig.
22). Wing brownish, paler than in female but darker along veins. Halter pale. Abdomen
dark brown; with abundant, semi-erect, long, single-striated scalelike hairs.
Genitalia (Fig. 26): Brown; proximal half of basistyle yellowish. Dististyle straight,
relatively stout, tapered to pointed tip. Aedeagus (Fig. 25) shorter and broader than
Wirth: Forcipomyia bicolor Group
72 20
\20
72 21
23
29 22
72
/ \
40 I
25 2
24 2
Figs. 20-26. Forcipomyia lacrimotorii; 20-21, 24, female; 22-26, male; from Santa
Catarina, Brazil (Plaumann): 20, antennal segments 8-15; 21, palpus; 22, fifth tarsomere
and claws of fore leg; 23, apex of hind tibia and basitarsus of hind leg; 24, spermathecae;
25, aedeagus; 26, genitalia, aedeagus and one dististyle omitted (scale in microns).
in bicolor and dubia, 1.4 times as long as basal breadth, basal arms and basal arch well
developed, anterior margin well sclerotized; sides slightly convex, tapering to blunt
apex; three well-marked ventral ridges at base. Parameres typical of bicolor Group;
bases well separated, distal portions straight, slender, tips slightly surpassing tip of
aedeagus.
Distribution.-Brazil.
Types.-Syntypes, 10 males, 13 females, Nova Teutonia, Santa Catarina, Brazil, F.
Plaumann, dates from 13.v.1937 to 7.ix.1938 (BMNH). Through the courtesy of Bruce
Townsend I have studied one male and two female syntypes from the British Museum
(Nat. Hist.). They are mounted on one slide with the following labels: "Brasilien / Nova
Teutonia / 270 11' B 52 23' L / Fritz Plaumann / 13.5.37." and "Forcipomyia / lacrimotorii
/ o o; SYNTYPE" in blue-bordered circular label and "PARATYPES" in red-bordered
blue label.
The two females are not dissected but mounted whole on their sides under one large
coverslip. One of these is in excellent condition; the other is damaged and has the
515
516 Florida Entomologist 74(4) December, 1991
antennae mounted separately under a small square coverslip. The male is mounted under
a separate small square coverslip and the genitalia and legs from one side have been
dissected off. The head is missing. This male is hereby designated LECTOTYPE.
I have also examined three male and three female pinned syntypes with data as
above. These are in moderately good condition, some with mesonotum somewhat rubbed
and vestiture missing. The characteristic mesonotal hair tufts are present on the better
specimens.
Other Specimens Examined.-Seven males, 10 females, same data as types but dates
viii.1945, v.1963, viii.1963, viii-ix.1970, ix.1970, i.1971, ii.1971 (USNM). Specimens from
this series have been selected as PLESIOTYPES and described and illustrated.
ACKNOWLEDGMENTS
I wish to express special appreciation to Maria Luiza Felippe-Bauer of the Instituto
Oswaldo Cruz in Rio de Janeiro, Brazil, and to Bruce C. Townsend of the Natural
History Museum, London, U.K. (BMNH) for the loan of types without which this study
would not have been possible. Most of the Brazilian specimens for this study were
purchased by the author from Fritz Plaumann of Nova Teutonia, Santa Catarina, Brazil,
many years ago and deposited in the National Museum of Natural History, Smithsonian
Institution, Washington, D.C. (USNM).
Financial support for this publication by Mars Incorporated, Effem Services Inc.,
Information Services International Division, Mt. Olive, NJ, is gratefully acknowledged.
REFERENCES CITED
BILLES, D. J. 1941. Pollination of Theobroma cacao L. in Trinidad, B.W.I. Trop.
Agric. (Trinidad) 18: 151-156.
BYSTRAK, P. G. AND W. W. WIRTH. 1978. The North American species of For-
cipomyia, subgenus Euprojoannisia (Diptera: Ceratopogonidae). U.S. Dept.
Agric. Tech. Bull. 159: 1-51.
DEBENHAM, M. L. 1987. The biting midge genus Forcipomyia (Diptera:
Ceratopogonidae) in the Australasian Region (exclusive of New Zealand) III. The
subgenera Forcipomyia s. s., and Lepidohelea. Invertebr. Taxon. 1: 167-199.
DOWNES, J. A., AND W. W. WIRTH. 1981. Ceratopogonidae, pp. 393-421, in McAlpine,
J. F., et al. [eds.], Manual of Nearctic Diptera, Vol. 1. Agric. Canada Monogr.
no. 27: 674 pp.
KAUFMANN, T. 1975a. Ecology and behavior of cocoa pollinating Ceratopogonidae in
Ghana, W., Africa. Environ. Entomol. 4: 347-351.
KAUFMANN, T. 1975b. Studies on the ecology and biology of a cocoa pollinator, For-
cipomyia squamipennis I. & M. (Diptera, Ceratopogonidae), in Ghana. Bull.
Entomol. Res. 65: 263-268.
LUTZ, A. 1914. Contribuicao para o conhecimento das Ceratopogoninas do Brazil.
Terceira memorial. Aditamento terceiro e descricao de species que nao sugam
sangue. Mem. Inst. Oswaldo Cruz 6: 81-99, 2 plates.
MACFIE, J.W.S. 1939. A report on a collection of Brazilian Ceratopogonidae (Dipt.).
Revta. Entomol. 10: 137-219.
MACFIE, J.W.S. 1944. Ceratopogonidae collected in Trinidad from cacao flowers. Bull.
Entomol. Res. 35: 297-300.
POSNETTE, A. F. 1950. The pollination of cacao in the Gold Coast. Hort. Sci. J. 45:
155-163.
SAUNDERS, L. G. 1956. Revision of the genus Forcipomyia based on characters of
all stages (Diptera, Ceratopogonidae). Canadian J. Zool. 34: 657-705.
SAUNDERS, L. G. 1959. Methods for studying Forcipomyia midges, with special ref-
erence to cacao-pollinating species (Diptera, Ceratopogonidae). Canadian J. Zool.
37: 33-51.
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 517
SORIA, S. DE J., AND W. W. WIRTH. 1974. Identidade e caracterizacao taxonomica
preliminary das mosquinhas Forcipomyia (Diptera, Ceratopogonidae) associadas
com a polinazacao do cacaueiro na Bahia. Revta. Theobroma 5(54): 3-22.
WINDER, J. A. 1977. Field observations on Ceratopogonidae and other Diptera:
Nematocera associated with cocoa flowers in Brazil. Bull. Entomol. Res. 67: 57-63.
WINDER, J. A. 1978. Cocoa flower Diptera: their identity, pollinating activity and
breeding sites. PANS 24: 5-18.
WIRTH, W. W. 1974. A catalogue of the Diptera of the Americas south of the United
States. 14. Family Ceratopogonidae. Mus. Zool. Univ. Sao Paulo 14: 1-89.
WIRTH W. W. 1991. New and little-known species of Forcipomyia (Diptera:
Ceratopogonidae) associated with cocoa pollination in Brazil. Proc. Entomol. Soc.
Washington. 93: 163-175.
WIRTH, W. W., N. C. RATANAWORABHAN, AND D. H. MESSERSMITH. 1977. Natural
History of Plummers Island, Maryland. XXII. Biting midges (Diptera:
Ceratopogonidae). 1. Introduction and key to genera. Proc. Biol. Soc. Washington
90: 615-647.
YOUNG, A. M. 1983. Seasonal differences in abundance and distribution of cocoa-pol-
linating midges in relation to flowering and fruit set between shaded and sunny
habitats of the La Lola cocoa farm in Costa Rica. J. Appl. Ecol. 20: 801-831.
aL -L a- a-- a a- a aaa
FIVE NEW SPECIES OF APHODIUS
(COLEOPTERA: SCARABAEIDAE)
FROM FLORIDA POCKET GOPHER BURROWS
PAUL E. SKELLEY
Department of Entomology & Nematology
Building 970, Hull Road
University of Florida
Gainesville, FL 32611-0740
ROBERT E. WOODRUFF
Florida State Collection of Arthropods
P. O. Box 147100
Gainesville, FL 32614-7100
ABSTRACT
Five new species of Aphodius (dyspistus, tanytarsus, hubbelli, platypleurus, and
pholetus), collected in the burrows of pocket gophers (Geomys pinetus Rafinesque), are
described. Similarities between Florida and Great Plains pocket gopher burrow faunas
are discussed, and reasons for the diversity ofAphodius in this habitat are postulated.
RESUME
Se described cinco nuevas species de Aphodius (dyspistus, tanytarsus, hubbelli,
platypleurus, y pholetus), colectadas en las galerias de Geomys pinetus Rafinesque. Se
discuten las similitudes entire la fauna de las galerias del genero Geomys de Florida y
de las Grandes Planicies. Se postulan las razones de la diversidad de Aphodius en estos
habitats.
Florida Entomologist 74(4)
Arthropods associated with animal nests and burrows are rarely studied with the
same vigor as their vertebrate hosts. This is especially true of burrowing animals and
their inquilines, where expensive or labor intensive sampling techniques are required.
Arthropods associated with the Southeastern pocket gopher, Geomys pinetus
Rafinesque, were studied by Hubbell & Goff (1939). Their work included accounts of
the rodent's habits, burrow structure, the associated arthropods, and a collecting
technique for burrow inhabiting arthropods that required minimal work and expense.
This method involved digging a hole to open the burrow system, removing the rodent
within, setting a baited pitfall trap in the exposed burrow, and then covering the hole
with a board and soil.
Hubbell & Goffs sampling method was used to collect some "rare" beetles reported
from this subterranean habitat: Geomysaprinus tibialis Ross (1940), Atholus minutus
Ross (1940), Spilodiscus floridanus Ross (1940) (Histeridae); Aphodius laevigatus Hal-
demann (1848) (= Aphodius goffi Cartwright 1939), and A. aegrotus Horn (1870a) (=
A. geomysi Cartwright 1939) (Scarabaeidae) (Woodruff 1973). Also collected were On-
thophilus kirmi Ross (1944) (not reported from Florida), 0. giganteus Helava (1978)
(previously known only from the holotype) (Histeridae), and 5 new species of the genus
Aphodius. Future sampling of pocket gopher burrows with this technique will undoub-
tedly produce new state records, rarely collected species, and other new species.
MATERIALS AND METHODS
Photographs were taken with a Hitachi S-450 scanning electron microscope (SEM).
Specimens were coated with 24k gold by an Eiko Engineering IB-2 Sputter Coater. The
dorsal habitus views of Aphodius dyspistus, n.sp., and Aphodius sepultus Cartwright
(1944) appear distorted because of their orientation.
The following abbreviations indicate collections where specimens are deposited:
FSCA Florida State Collection of Arthropods; GHNC G. H. Nelson Collection; HAHC
H. F. Howden Collection; PESC P. E. Skelley Collection; PMCC P. M. Choate
Collection; PWKC P. W. Kovarik Collection; REWC R. E. Woodruff Collection;
RHTC R. H. Turnbow Collection; RWLC R. W. Lundgren Collection; WBWC W.
B. Warner Collection; USNM United States National Museum of Natural History.
Aphodius dyspistus Skelley & Woodruff, n.sp.
(Fig. 1-2, 5-6)
HOLOTYPE: Male; length 3.2mm, width 1.5mm. Body elongate, dorso-ventrally
flattened, alutaceous, feebly shiny and roughly sculptured dorsally; dorsal punctures
coarse, setiferous. Color dark brown; legs, mouthparts, antennae and clypeus yellowish-
brown.
Head convex with 2 weak tubercles, surface granulate, alutaceous between granules;
punctures setiferous and indistinct, except at base of head; clypeus slightly emarginate,
broadly rounded and reflexed on each side; genae and clypeus form a slight angle, visible
in lateral view. Genae with evenly rounded fimbriate margin.
Pronotum convex, length 0.75 of width, sides arcuate, narrower in front; hind angles
well defined, obtuse, slightly reflexed; front angles nearly 90, slightly reflexed; base
moderately sinuate, fine margin obscured by punctures; disc with dense coarse setiferous
punctures; punctures of 2 sizes, smaller punctures half diameter of larger; setae recurved,
short, not or barely reaching edge of neighboring punctures.
Elytra deeply striate, striae as wide or wider than intervals; strial punctures coarse,
apparently in pairs, with a transverse ridge connecting each pair, giving striae a seg-
mented or pitted appearance, strial punctures without setae; intervals convex, with
December, 1991
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 519
Fig. 1-2. Aphodius dyspistus n.sp., male genitalia, line = 0.lmm. 1.) Right lateral
view. 2.) Dorsal view.
double row of punctures bearing short recurved setae, not reaching neighboring
punctures.
Mesosternum alutaceous, slightly convex between coxae. Metasternum shining, sides
alutaceous, medial area flattened, weakly alutaceous, with fine scattered punctures.
Abdominal segments alutaceous with scattered setiferous punctures, setae nearly as
long as the segment.
Protibia laterally tridentate, serrate above teeth; apical spur acute, slightly curved
ventrally, as long as tarsal segments 1-2 combined, not surpassing apical tooth. Profemur
alutaceous, with coarse pictures on posterior surface. Meso- and metatibia with an
apical fringe of unequal length, longest setae equal in length to short spur. Short spur
of mesotibia truncate at tip, half length of long spur, with a fine tooth on medial apex;
long inner spur slightly curved ventrally. Short metatibial spur 0.75 length of long spur,
both straight. Meso- and metafemora twice as long as wide, weakly alutaceous, with
scattered setiferous punctures.
Protarsal 1st segment half length of tibial spur, segments 1-4 nearly equal in length,
5th segment equal to 3+4 combined, claw length equal to 4th segment. Meso- and
metatarsal 1st segment slightly longer than longest tibial spur, equal to 2 + 3 combined;
segments 2-4 equal in length, 5th 0.75 length of 3+4 combined, claws equal in length
to 4th segment; segments 1-4 with a double ventral row of setae, stoutest and most
numerous on 1st segment.
Genitalia as in Figs. 1-2.
Florida Entomologist 74(4)
Fig. 3-4. Aphodius sepultus Cartwright. 3.) Elytral striae near scutellum, line =
0.1mm. 4.) Dorsal view, line = 0.5mm.
ALLOTYPE: Female; length 3.5mm, width 1.7mm. Similar to holotype except short
metatibial spur acute, not truncate or toothed.
Variations: Length 2.9-3.5mm, width 1.4-1.7mm. Some specimens are more reddish-
brown than others.
Specimens examined: Holotype and Allotype: FLORIDA, ALACHUA CO., 2.5 mi.
SW of Archer, 5-11-1-1990, P. E. Skelley, Geomys burrow pitfall (FSCA). Paratypes
(36 Males, 71 Females): FLORIDA, ALACHUA CO., 2.5 mi. SW of Archer, 16-1-1988,
P. E. Skelley, Geomys dung chamber (6M,10F: REWC, PESC); loc. cit., 11-18-1-1988,
P. E. Skelley, window trap in old pasture (1F: PESC); loc. cit., various dates 7-I to
24-III-1988, P. E. Skelley, Geomys burrow pitfall (11M,27F: GHNC, HAHC, PWKC,
REWC, RHTC, USNM); loc. cit., various dates 8-I to 17-II-1989, P. E. Skelley, Geomys
burrow pitfall (2M,3F: USNM); loc. cit., various dates 27-XII-1989 to 15-1-1990, P. E.
Skelley, Geomys burrow pitfall (11M,17F: USNM, WBWC); loc. cit, 20-1-1991, P. E.
Skelley, R. H. Turnbow, F. Skillman, Geomys dung chambers (1M,5F: FSCA); Gaines-
ville, Doyle Conner Building, various dates 4-II-24-III-1988, P. E. Skelley, Geomys
burrow pitfall (1M,5F: PESC); 4 mi. W of 1-75 on Rt. 24, 17-IV-1983, P. M. Choate,
Geomys burrow (2M: PMCC); GILCHRIST CO., nr. Trenton, 2.3 mi. W of Alachua Co.
line on St. Rd. 26, 7-15-1-1989, P. E. Skelley, Geomys burrow pitfall (1F: REWC);
LEVY CO., 2930'N, 82035'W, 4.0 mi. SW of Archer, 29-1-1990, R. W. Lundgren,
Geomys burrow pitfall (2F: RWLC); WALTON CO., Defuniak Springs Airport, 14-21-I-
1990, P. E. Skelley, R. H. Turnbow, & M. C. Thomas, Geomys burrow pitfall (1M,2F:
REWC, RHTC); loc. cit., 28-II to 6-III-1990, R. H. Turnbow, Geomys burrow pitfall
(1M: RHTC).
December, 1991
520
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 521
Fig. 5-6. Aphodius dyspistus n.sp. 5.) Elytral striae near scutellum, line = 0.1mm.
6.) Dorsal view, line = 0.5mm.
Remarks: Aphodius dyspistus specimens are often covered with dirt and require
cleaning with a wet brush before surface structures can be seen.
Relationship: This species falls into Horn's (1887) group H, and is similar to A.
sepultus Cartwright (1944), in both morphology and association with pocket gophers.
These species are readily separated by the following characters: 1) long, recurved pro-
notal and elytral setae reaching well into neighboring punctures in sepultus; shorter
setae not or barely reaching neighboring punctures in dyspistus; 2) flattened elytral
intervals wider than striae in sepultus; convex intervals narrower than striae in dyspis-
tus; 3) an "unusual twinning or doubling of the elytral stria" (Cartwright 1944) with
indistinct punctures in sepultus; coarse, paired strial punctures giving striae a segmented
or pitted appearance in dyspistus (Fig. 3-6).
Etymology: Greek; dys = hard, with difficulty; pistos = trusted, genuine; dys-pistos
= hard to believe, incredible. Named for the surprise that this and the following species
have remained unknown, considering the fieldwork previously done in Florida.
Aphodius tanytarsus Skelley & Woodruff, n.sp.
(Fig. 7, 8, 18-19)
HOLOTYPE: Male; length 4.4mm, width 1.9mm. Body elongate, slightly dorso-ven-
trally flattened, shiny throughout, dorsal punctures without setae. Color dark brown
to black; apical clypeal margin, anterior pronotal angles, legs, mouthparts, antennae,
and elytra reddish brown.
Florida Entomologist 74(4) December, 1991
'K
Fig. 7. Dorsal habitus Aphodius tanytarsus n.sp., line = 2.0mm.
522
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 523
Fig. 8. Aphodius tanytarsus n.sp., mesotibia and tarsus, line = 0.5mm.
Head surface convex, trituberculate; base punctate, size intermediate between fine
and coarse pronotal punctures; clypeus granulate, shallowly emarginate, broadly
rounded, reflexed on each side. Gena and clypeus form distinct angle where they meet,
visible in lateral view. Genae fimbriate, distinct on dorsal view, slightly angled at post-
erior third.
Pronotum convex, length 0.70 width, sides arcuate, slightly narrower in front; hind
angles obtuse; front angles nearly 900, slightly reflexed; base evenly rounded, margin
complete and distinct. Disc with regularly placed fine punctures and irregularly placed
coarse punctures, 5 times diameter of fine ones; coarse ones not present medially at
pronotal sides.
Elytra broadest medially, shiny; striae fine and distinct, strial punctures separated
by twice their diameters, connected by a continuous groove; intervals flattened with 2
irregular rows of minute punctures.
Mesosternum alutaceous, not carinate between coxae. Metasternum alutaceous ex-
cept for flattened shiny median area, entire surface finely punctate, punctures of alutace-
ous sides setiferous. Abdominal segments alutaceous with scattered setiferous punctures,
setae as long as the segments.
Protibia laterally tridentate, slightly serrate above teeth; spur acute, curved vent-
rally, not surpassing apical tooth. Profemur alutaceous with scattered punctures on
posterior surface. Meso- and metatibiae with apical fringe of unequal length, long fringe
setae longer than short spur. Short mesotibial spur half length of long spur, bluntly
rounded at tip, with small tooth on median apex. Short metatibial spur 0.75 length of
long spur, both straight, acute. Meso- and metafemora with few fine setiferous punctures.
Mesofemur with coarse punctures bearing long setae on apical third.
Protarsal segments 1-4 equal in length; 5th segment, claws, and segments 3+4
combined equal in length to the protibial spur. Meso- and metatarsal 1st segment, 5th
segment, and claws equal in length to long tibial spur (fig. 8); 2nd segment half length
of 1st; segments 2-4 equal in length; segments 1-4 with a double ventral row of stiff
setae, stoutest and most numerous on 1st segment. Mesotarsus longer than mesotibia.
Genitalia as in Figs. 18-19.
ALLOTYPE: Female; length 4.5mm, width 2.0mm. Similar to holotype except outer
spur of mesotibia acute, not truncate or toothed.
Variations: Length 4.2-4.7mm, width 1.8-2.1mm. The coarse pronotal punctures of
some specimens are denser at the pronotal angles, occasionally coalescing.
Specimens examined: Holotype and Allotype: FLORIDA, ALACHUA CO., 2.5 mi.
SW of Archer, 5-11-1-1990, P. E. Skelley, Geomys burrow pitfall (FSCA). Paratypes
(17 Males, 13 Females): FLORIDA, ALACHUA CO., 2.5 mi. SW of Archer, 16-1-1988,
P. E. Skelley, Geomys dung chamber (2M: REWC); loc. cit., various dates 23-I to
3-II-1988, P. E. Skelley, Geomys burrow pitfall (2M,1F: REWC); loc. cit., various dates
2-11-1-1990, P. E. Skelley, Geomys burrow pitfall (2M,2F: PESC); loc. cit, 20-1-1991,
524 Florida Entomologist 74(4) December, 1991
P. E. Skelley, R. H. Turnbow, F. Skillman, Geomys dung chambers (3M: FSCA);
OKALOOSA CO., 2.8 mi. W of Walton Co. line on US-90, Deerland, various dates
3-21-1-1990, P. E. Skelley, P. W. Kovarik, R. H. Turnbow, & M. C. Thomas, Geomys
burrow pitfall (1M,4F: GHNC, HAHC, PESC, PWKC); WALTON CO., Defuniak
Springs Airport, various dates 14-26-1-1990, P. E. Skelley, R. H. Turnbow, & M. C.
Thomas, Geomys burrow pitfall (6M,6F: HAHC, REWC, RHTC, USNM, WBWC); loc.
cit., 6-12-III-1990, R. H. Turbow, Geomys burrow pitfall (1M: RHTC).
Remarks: Aphodius tanytarsus falls into Horn's (1887) group I-d, but is not similar
to any known species. The long tarsal claws, as long as the long meso- and metatibiae
spurs (Fig. 7-8), distinguish this species from any other seen.
Etymology: Greek; tany = stretched; tarsus = foot; tany-tarsus = stretched foot.
The name refers to its unusually long, tarsal claws.
Aphodius hubbelli Skelley & Woodruff, n.sp.
(Fig. 10, 14-15, 16-17, 20-21)
Aphodius haldemani Horn 1887 in Woodruff 1973:92; Woodruff 1982:90.
HOLOTYPE: Male; length 8.8mm, width 3.8mm. Body elongate, dorso-ventrally
flattened, shiny throughout. Body and appendages reddish-brown.
Head surface convex, smooth, shiny, minute punctures at base. Clypeus shallowly
emarginate, broadly rounded on each side, slightly reflexed. Genae indistinct in dorsal
view, fimbriate.
Pronotum convex, length 0.625 width, lateral margin arcuate, narrower at front;
sides strongly reflexed with coarse punctures evenly distributed (Fig. 10), separated by
distance equal to their diameters; hind angles broadly rounded, anterior angles more
acutely rounded; pronotal disc impunctate, shiny; base sinuate, not margined.
Elytra broadest medially, shiny; striae fine, punctures separated by distance equal
to their diameters; intervals flat, impunctate on disc, minutely punctate laterally.
Mesosternum alutaceous, not carinate between coxae. Metasternum alutaceous with
coarse setiferous punctures laterally, shiny with minute punctures medially. Abdominal
segments alutaceous, with scattered setiferous punctures, setae half as long as the
segment.
Protibia laterally tridentate, with a dense row of ventrally projecting setae on medial
edge; spur spatulate, parallel sided and truncate at the tip (Fig. 14), curved ventrally,
surpassing apical tibial tooth. Profemur with a patch of long dense setae on ventral
margin, scattered setiferous punctures on posterior surface. Mesotibia with apical setal
fringe of unequal length, long setae equal in length to short spur. Short spur half length
of long spur, both straight, narrow and acute. Mesotibia with medial patch of dense
setae on apical 0.80. Mesofemur with coarse punctures near apex; postero-ventral margin
with a dense patch of setae on medial 0.60. Meso- and metatrochanter with 3-4 setae.
Metafemur with a patch of setae on posterior margin near the trochanter, 0.40 times
length of femur; coarse punctures present near apex. Metatibial apical fringe half length
of short spur. Short spur 0.75 length of long spur, both straight, narrow and acute.
Metatibia with lateral surface flattened, ventral edge with subapical process, medially
with a patch of setae from tooth to apex (Fig. 16-17).
Protarsi half length of protibia, segments 1-4 equal in length, 1-3 together equal
length of spur, 5th segment equals 3+4 combined, claws 0.75 length of 5th segment.
Meso- and metatarsi same length as their corresponding tibia, proportions similar; seg-
ments 2-4 decreasing in length; 2nd segment 0.50 length of 1st, 4th segment 0.75 length
of 2nd, 5th segment 0.625 length of 1st, claws 0.75 length of 5th. Meso- and metatarsi
differ in mesotibial long spur equal in length to 1st tarsal segment and the metatibial
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 525
a
Fig. 9-11. Pronotum, left lateral view, line = 1.0mm. 9.) Aphodius magnificens
Robinson. 10.) Aphodius hubbelli n.sp. 11.) Aphodius haldemani Horn.
Florida Entomologist 74(4)
Fig. 12-15. Dorsal view of left protibial spur, line = 0.5mm. 12.) Aphodius magnifi-
cens Robinson, male. 13.) Aphodius haldemani Horn, male. 14.) Aphodius hubbelli
n.sp., male. 15.) Aphodius hubbelli n.sp., female.
long spur 0.66 length of 1st tarsal segment. Meso- and metatarsal segments 1-4 with
double ventral row of stiff setae, stoutest and most numerous on 1st segment.
Genitalia as in Figs. 20-21.
ALLOTYPE: Female; length 8.4mm, width 3.7mm. Similar to holotype except pro-
tibial spur curved medially and pointed, not spatulate (Fig. 15), barely surpassing apical
tibial tooth; length of protibial spur equals length of 1st and 2nd tarsal segments combined;
and dense patches of setae on tibia, femur, and trochanter lacking.
Variation: Length 8.1-9.3mm.; width 3.5-4.2mm. Pronotal punctures vary in number,
but are always present and evenly distributed along pronotal side. Male metatibial
characters vary in size of subapical process, and in number of punctures on lateral surface.
526
December, 1991
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 527
16
Fig. 16-17. Left metatibia of male Aphodius hubbelli n.sp., line = 1.0mm. 16.)
Lateral view. 17.) Medial view.
Specimens Examined: Holotype and Allotype: FLORIDA, OKALOOSA Co., 2.8 mi.
W of Walton Co. line on US-90, Deerland, 3-13-1- 1990, P. E. Skelley & P. W. Kovarik,
Geomys burrow pitfall (FSCA). Paratypes (8 Males, 5 Females): FLORIDA: ALACHUA
CO., 2.5 mi. SW of Archer, 8-22-1-1989, P. E. Skelley, Geomys burrow pitfall (2F:
PESC); Gainesville, 9-1-1969, R. E. Woodruff & D. L. Mays, malt pitfall (1M: REWC);
4 mi. W of 1-75 on Rt. 24, 17-IV-1983, Geomys burrow, P. M. Choate (1M,2F: PMCC);
OKALOOSA CO., 2.8 mi. W of Walton Co. line on US-90, Deerland, various dates
3-26-1-1990, P. E. Skelley, P. W. Kovarik, R. H. Turnbow & M. C. Thomas, Geomys
burrow pitfall (3M: HAHC, PESC, USNM); WALTON CO., Defuniak Springs Airport;
13-21-1-1990, P. E. Skelley, R. H. Turnbow & M. C. Thomas, Geomys burrow pitfall
(3M,1F: USNM, REWC, RHTC).
Remarks: This species keys to A. haldemani Horn (1887) in both Horn (1887, group
I-a) and Woodruff (1973). Woodruff (1973:92, fig. 180; 1982:90) described and illustrated
the single specimen from Gainesville (mentioned above) under the name A. haldemani,
stating it was possibly a new species. Additional specimens have confirmed it as distinct.
Aphodius hubbelli is closely related to A. magnificens Robinson (1938) and A. halde-
mani. Males of A. hubbelli can be distinguished by their truncate parallel-sided protibial
spur and the presence of the subapical process on the metatibia. Aphodius magnificens
and A. haldemani have the protibial spurs apically divergent, triangular in shape (Fig.
12-14) and lack the metatibial process. Both sexes of these 3 species can be separated
by the distribution and spacing of the pronotal punctures (Fig. 9-11). Aphodius hubbelli
has these punctures evenly distributed and spaced.
Etymology: This species is named in honor of the late T. H. Hubbell for his many
contributions to Florida biogeography (Hubbell 1954, 1961; Hubbell et al. 1956) and his
pioneering studies (with Goff 1939) on the arthropods of pocket gopher burrows, including
the description of a new genus (Typhloceuthophilus) of blind camel-crickets (Hubbell
1940).
*
;7~--+4~
Florida Entomologist 74(4)
I I
22
23
Fig. 18-25. Genitalia of males, lines = 0.5mm. 18-19.) Aphodius tanytarsus n.sp.
18.) Lateral view. 19.) Dorsal view. 20-21.) Aphodius hubbelli n.sp. 20). Lateral views.
21.) Dorsal view. 22-23). Aphodius platypleurus n.sp. 22.) Lateral view. 23.) Dorsal
view. 24-25). Aphodius pholetus n.sp. 24.) Lateral view. 25.) Dorsal view.
Aphodius platypleurus Skelley & Woodruff, n.sp.
(Fig. 23-23, 26)
HOLOTYPE: Male; length 7.4mm, width 3.5mm. Body elongate, parallel sided,
dorso-ventrally flattened, shiny throughout. Color dark chocolate-brown; elytra, legs,
and mouthparts paler; antennae and tarsi yellowish-brown.
Head convex, surface shiny, slightly alutaceous, with minute punctures separated
by 2-3 times their diameters. Clypeus slightly emarginate, narrowly rounded at sides,
reflexed. Genae moderately prominent, fimbriate. Gena and clypeus form a distinct
angle, visible in lateral view.
Pronotum convex, margined basally at medial 0.66, length 0.66 width; lateral margins
parallel except at anterior angles, margin thickest medially; sides broadly explanate and
flattened, shelf-like (Fig. 26), alutaceous with a few coarse punctures; all pronotal angles
equally rounded; disc with coarse punctures separated by 0-4 times their diameter,
absent on anterior fourth, coarse punctures 5 times larger than minute punctures; minute
punctures evenly scattered on disc.
Elytra weakly alutaceous, shiny; intervals flat. Striae groove-like; punctures, where
visible, connected by this groove.
December, 1991
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 529
'- -
/
j,
r4 Br
Fig. 26. Dorsal habitus of Aphodius platypleurus n.sp., line = 3.0mm.
Florida Entomologist 74(4)
Mesosternum coarsely punctate and alutaceous. Metasternum alutaceous throughout;
coarsely punctate laterally, punctures with minute setae. Abdomen alutaceous, with
indistinct coarse punctures bearing minute setae.
Protibia laterally tridentate, proximal tooth smallest, spur with apical third bent
ventro-medially, surpassing apical tibial tooth. Profemur alutaceous with a few irregular
longitudinal lines and scattered punctures on posterior surface, coarsely punctate with
setae on anterior surface. Meso- and metatibia with apical setal fringe of unequal length,
long setae half length of short spur.
Meso- and metatibial spurs narrow, pointed, slightly curved. Short spur of mesotibia
0.66 length of, and slightly broader than, long spur; medial mesotibial surface with
numerous setae as long as long apical fringe setae. Meso- and metafemora weakly
alutaceous. Mesofemur with coarse punctures near apex; posterior edge with a patch
of short, stout setae covering medial half. Metatibia with short spur 0.75 length of long
spur. Metafemur with coarse pictures near apex and small patch of short setae near
trochanter.
Protarsal segments 1-4 equal in length, segments 1-3 combined length equal to spur,
5th segment equals 3+4 combined. Meso- and metatarsal segments 2-4 subequal in
length, each segment half length of 1st; 5th segment 0.75 length of 1st, claws half length
of 5th segment; 1st segment with double ventral row of setae, row indistinct on other
segments. Mesotarsal 1st segment equals length of long spur, tarsi longer than mesotibia.
Metatarsal 1st segment 0.75 length of long spur, tarsi same length as metatibia.
Genitalia as in Figs. 22-23.
ALLOTYPE: Female; length 7.2mm, width 3.4mm. Similar to holotype except it
lacks the patches of setae on the meso- metafemur, and metatibia; and the mesotibial
short spur is the same width as the long spur.
Variation: Length 6.5-7.5mm, width 3.4-3.8mm. The punctures and alutaceous micro-
sculpture of the head and pronotum vary in their distinctness and density. The relative
thickness of the lateral pronotal margin also varies between specimens. Most have the
anterior half of the pronotal margin thicker than the posterior half, with the middle
being the thickest.
The 2 specimens from Okaloosa Co. are lighter in color, less shining, and have the
pronotal punctures sparser and less distinct than the type series. Because of this differ-
ence and the lack of a larger series, these 2 specimens are not designated as paratypes.
Specimens Examined: Holotype and Allotype: FLORIDA: LEVY CO., 3.0 mi. SW
of Archer, 29-1-1990, R. W. Lundgren, Geomys burrow pitfall (FSCA). Paratypes (4
Males, 8 Females): same data as holotype (4M,5F: HAHC, PESC, REWC, RWLC,
USNM); loc. cit., 25-1-1990 (1F: PESC); loc. cit., 9-II-1990 (1F: REWC); 4.0 mi. SW of
Archer, 15-22-XII-1990, P. E. Skelley, Geomys burrow pitfall (1F: PESC). Non-
Paratypes: OKALOOSA CO., 2.8 mi. W of Walton Co. line on US-90, Deerland, various
dates 3-21-1-1990, P. E. Skelley, P. W. Kovarik, R. H. Turnbow, & M. C. Thomas,
Geomys burrow pitfall (1M,1F: PESC).
Remarks: Aphodius platypleurus keys to A. brevicollis LeConte (1878) in Horn (1887
group I-a). Apparently it is related to A. russeus Brown (1928) and A. acuminatus
Cartwright (1944), but differs in having the pronotum greatly explanate with parallel
lateral margins, and a dark brown color. Aphodius russeus has the lateral pronotal
margins parallel, but not distinctly explanate, and A. acuminatus has the pronotal
margins explanate, but not parallel sided. Both are red or reddish-brown.
Etymology: Greek; platy = broad, flat; pleura = side; platypleurus = broad flat
sides. Named in reference to its greatly expanded and flattened pronotal sides.
December, 1991
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 531
Aphodius pholetus Skelley & Woodruff, n.sp.
(Fig. 24-25)
HOLOTYPE: Male; length = 6.5mm; width = 3.2mm. Body elongate, dorso-vent-
rally flattened, shiny, not alutaceous. Color chocolate-brown; elytra, lateral margins of
pronotum and head, mouthparts, antennae, legs, and abdomen paler.
Head convex, surface granulate on clypeus at median apical margin; otherwise
smooth, shiny, and impunctate. Clypeus slightly emarginate, broadly rounded on each
side, slightly reflexed. Slight angle formed between clypeus and gena (visible in lateral
view), gena otherwise indistinct. Posterior margin of gena forming 90 angle, with setae
at angle.
Pronotum convex, length 0.66 width; sides arcuate, anteriorly convergent from hind
angles; angles rounded, obtuse; sides moderately reflexed, without depressed areas;
base evenly arcuate, not sinuate or margined; disc impunctate anterio-medially; coarsely
punctate laterally and at base, punctures separated by their diameter, not or rarely
touching.
Elytral striae distinct, punctures separated by distance equal to their diameter,
connected by strial groove. Intervals slightly convex, dorsally impunctate; laterally with
fine punctures, becoming coarser at apex.
Mesosternum alutaceous, slightly convex between coxae, coarsely punctate laterally.
Metasternum alutaceous with coarse seta bearing punctures laterally; medially shiny
with minute punctures. Abdomen slightly alutaceous with setiferous punctures, setal
length about 0.20 segment length.
Protibia laterally tridentate, barely serrate above teeth; spur stout, curved ventrally,
acute, tip not surpassing apical tooth, length equal to protarsal segments 1-2 combined.
Profemur alutaceous and setiferous on anterior surface; slightly alutaceous with scattered
punctures on posterior surface.
Meso- and metatibia with apical fringe setae of unequal length. Mesotibial spur length
equal to length of long fringe setae, half length of long spur, apically blunt; apex slightly
twisted, tooth medial. Long mesotibial spur same width as short spur, narrow and acute.
Meso- and metatibial femora shiny, with 5-6 coarse punctures near apex and scattered
minute punctures on ventral surface, with sparse scattered setiferous punctures on
posterior edge. Meso- and metatrochanter each with a seta. Short metatibial spur 2
times length of long fringe setae, 0.75 length of long spur; both spurs narrow and acute.
Protarsal segments 1-4 equal in length, segment 1+2 length equal to spur, segment
5 to 3 +4 combined, claws 0.625 length of segment 5.
Mesotarsi equal to mesotibial in length, 1st segment equal in length to long spur,
2nd segment half length of 1st, 2-4 decreasing in length, 4th segment 0.75 length of
2nd, 5th segment 0.625 length of 1st, claws half length of 5th; segments 1-4 with double
ventral row of setae, stoutest and most numerous on 1st. Metatarsi same except 2nd
segment 0.375 length of 1st.
Genitalia as in Figs. 24-25.
ALLOTYPE: Female unknown.
Specimens examined: Known only from the holotype: FLORIDA: OKALOOSA CO.,
2.8 mi. W of Walton Co. line on US-90, Deerland, 4-1-1990, P. W. Kovarik & P. E.
Skelley, Geomys burrow (FSCA).
Remarks: Aphodius pholetus keys to Horn's (1887) group I-a and is closely related
to A. atwateri Cartwright (1944) and A. oklahomensis Brown (1928) but is distinct in
several characters. Aphodius pholetus lacks minute punctures on the head, pronotum,
Florida Entomologist 74(4)
and intervals of the elytral disc; has anteriorly convergent lateral pronotal margins; has
the coarse pronotal punctures well separated at the posterior angles; has the genae
more angular, not rounded, nearly 900, and lacks the depressions at the hind angles of
the pronotum. Both A. atwateri and A. oklahomensis have minute punctures on the
head, pronotum, and elytral disc; lateral pronotal margins parallel at the basal half;
coarse pronotal punctures coalescing at the posterior angles; the genae more rounded
with an angle greater than 900; and have depressions at the hind angles of the pronotum.
Etymology: Greek; pholeter = one who lurks in a hole. The type specimen was hand
collected at the bottom of a hole exposing a pocket gopher burrow.
PLACEMENT IN EXISTING KEYS
The following are modifications of Gordon's (1983) key to the Eastern North American
species of Aphodius, to include these species.
Elytra with deep wide striae, narrower convex intervals, pocket gopher burrows;
Florida ................................................................ dyspistus Skelley & Woodruff
- Elytra with fine narrow striae, wider flat intervals ........................... 12a
12a. Basal marginal line of pronotum strongly impressed; Georgia ..........
........................................ ........................................ fordi G ordon
- Basal marginal line of pronotum obsolete; not known from Georgia or
Florida (introduced European speices) ................................... scrofa (F.)
31. Tarsal claws same length as longtibial spurs; pocket gopher burrows;
Florida ................................................. tanytarsus Skelley & Woodruff
- Tarsal claws less than 0.75 length of long tibial spurs ....................... 31a
31a. Foretibia with apical spur long, abruptly hooked and acuminate, or
broadly expanded, spatulate; pocket gopher burrows .......................... 32
- Foretibia with apical spur variably modified, but never as described
above; various habitats ............................................. ............... 33
32. Disc of pronotum smooth, not punctate medially; pronotal lateral margins
explanate; reddish brown in color ................................................... 32a
- Disc ofpronofum punctate medially, with 2 puncture sizes; pronotal lateral
margins explanate or not ............................................................. 32b
32a. Coarse lateral punctures of pronotum evenly distributed; male protibial
spur parallel sided and truncate; male metatibia with a subapical tooth
on ventral edge; pocket gopher burrows; Florida ...........................
.................. ..................................... hubbelli Skelley & W oodruff
Coarse lateral punctures of pronotum not evenly distributed, nearly coa-
lescing on posterior angles; male protibial spur triangular; male metatibia
without subapical tooth; the Great Plains ....................... haldemani Horn
32b. Pronotum with lateral margin slightly explanate; Iowa and the Great
Plains ......................................................................... russeus Brown
Pronotum with lateral margins greatly explanate, flattened, parallel
sided; pocket gopher burrows; Florida ..... platypleurus Skelley & Woodruff
41. Lateral margin of pronotum explanate; elytra entirely brown; rodent
burrow s or nests .......................................................................... 41a
Lateral margins of pronotum not explanate, or feebly so; elytra black or
brown, or variably marked; not konwn to occur in rodent burrows ........ 42
41a. Pronotum with evenly distributed fine and coarse punctures; western
Canada ................................................................... leptotarsis Brown
532
December, 1991
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 533
- Pronotum without fine punctures; coarse punctures restricted to the base
and sides of pronotum; pocket gopher burrows; Florida ................
...I.................................................... .... pholetus Skelley & W oodruff
The following are modifications of Woodruff's (1973) key to the Aphodius of Florida.
3(2'). Clypeus one each side angulate; elytral pubescence short, inconspicuous,
scattered; length 5.5-7mm ......................................... lutulentus Hald.
3'. Clypeus on each side broadly rounded, barely angulate; elytral pubes-
cence obvious; length 3-5 mm ..................................... ............. 3a
3a(3'). Pronotal punctures coarse, nearly coalescing, with setae large and prom-
inent; pocket gopher burrows ..................... dyspistus Skelley & Woodruff
3a'. Pronotal punctures separated by distance equal to their diameter, not
coalescing, without visible setae ...................................... stupidus Horn
4(1'). Pronotum broadly explanate, flattened and parallel sided; color dark
chocolate-brown, elytra often paler; pocket gopher burrows ...................
......................................................... platypleurus Skelley & W oodruff
4'. Pronotum not explanate at sides; or if so, then the pronotal sides are
evenly curved, not parallel sided ..................................... .......... 4a
4a(4'). Size larger (length 6-8mm; width 3-4mm); elytra red or brown, not black 5
4a'. Size smaller (length 2-6.5mm; width 1-2.5mm); elytral color variable from
black to pale yellow ....................................................................... 6
5(4a). Bicolored, elytra red, the pronotum black with anterior angles red. Head
with three prominent tubercles; common species in cow dung fimetarius (L.)
5'. Not as distinctly bicolored, elytra at most slightly paler; head without
tubercles; pocket gopher burrows, not known from cow dung ............. 5a
5a(5'). Color dark brown, elytra paler .................... pholetus Skelley & Woodruff
5a'. Color red or reddish-brown ........................................... ........... 19
9(8'). Tarsal claws as long as long tibial spurs; pocket gopher burrows ........
...................................................... ..... tanytarsus Skelley & W oodruff
9'. Tarsal claws less than 0.75 length of long tibial spurs ....................... 9a
9a(9'). Base color of elytra and pronotum yellow with smoky brown markings
................................................................................. lividus (Oliv.)
9a'. Base color black to dark red brown, nearly unicolorus ........................ 10
19(5a'). Male anterior tibial spur broad and spatulate shaped; pronotum explanate
at sides, pocket gopher burrows (replaces A. haldemani in original
key) ....................................................... hubbelli Skelley & Woodruff
19'. Anterior tibial spur normal, narrow and pointed; pronotum normal, not
explanate at sides; pocket gopher burrows and locally frequent at light
.............................................................................. laevigatus H ald.
DISCUSSION
During our search for Aphodius species related to these new species, similarities
between the Coleoptera faunas of pocket gopher burrows in the Great Plains and Florida
became apparent (Blume & Aga 1975, 1979, Blume & Summerlin 1988). Some species
are found in both regions; Onthophilus kirni Ross (1944) and Geomysaprinus goffi Ross
(1940) (Histeridae). Others are closely related or sister species: Spilodiscus floridanus
Ross (1940) and S. gloveri (Horn 1870b) (Histeridae); Aphodius laevigatus Haldeman
(1848) and A. concavus Say (1823); A. dyspistus n.sp. and A. sepultus Cartwright (1944);
Florida Entomologist 74(4)
A. tanytarsus n.sp. and A. insolitus Brown (1928); A. hubbelli n.sp. and A. haldemani
Horn (1887); A. platypleurus n.sp. and A. acuminatus Cartwright (1944); A. pholetus
n.sp. and A. atwateri Cartwright (1944). Other burrow inhabiting taxa (Staphylinidae,
Leptodiridae, Gryllacrididae (Orthoptera), Diptera, Collembola, etc.) need considerable
work before any corresponding similarities might be discovered. Before the taxonomy
of these poorly known groups can be settled and overall similarities in their faunas
elucidated, far more collecting is needed throughout the ranges of geomyid rodents.
Many questions about partitioning and species competition arise when several species
of Aphodius are collected from pocket gopher burrows at one locality. Such species
diversity is striking in such a small ecological niche.
Although we have no specific observations, some hypotheses about niche partitioning
in this habitat include: 1.) differences in soil conditions, moisture, temperature, ventila-
tion, composition, etc. within the same burrow system; 2.) differences in gross habitat
conditions, such as old field vs. scrubby woods; 3.) the specific food source of the rodent;
e.g. roots, stems, or greens of various plants and their availability; 4.) burrow variations
due to differing behaviors of the sexes, possibly correlated with 5.) the age of the burrow
system. 6.) All burrow systems are similarly constructed with tunnels connecting food,
fecal, and nest chambers. Pocket gopher dung (the presumed larval food) is normally
deposited in fecal chambers, but some is scattered in the burrow system. Larger species
(e.g. A. laevigatus and A. hubbelli) may require more than a single pellet for larval
development and may be restricted to fecal chambers. Smaller species (e.g. A. dyspistus)
could complete larval development in a single pellet and adapt to those scattered along
the tunnel. 7.) Perhaps A. aegrotus and A. laevigatus, the 2 most common species in
the burrow, are partially temporally isolated from the other rarer species. The rarity
of the other species may be partially accounted for by a.) their activity during the colder
months, b.) a lower reproductive potential, c.) the utilization of the food source by
A. aegrotus and A. laevigatus during the major portion of the year, or any combination
of the above.
Pocket gopher burrows were surveyed for an entire year by using the method de-
scribed by Hubbell & Goff (1939). This capture data, in combination with the light trap
collection data reported by Woodruff (1973), indicates activity periods for the Aphodius
fauna of the burrows. Aphodius laevigatus and A. aegrotus were collected from the
burrows and at light throughout the year. The majority of A. aegrotus captures were
in fall (August to November) and in the winter (January to February) for A. laevigatus.
The 5 new species have not been taken at light. Aphodius dyspistus has been captured
in the borrows from December to April. The other 4 species have been captured, almost
exclusively, in January.
In order to disperse, beetles must either find connecting burrows or exit to the
surface and find another burrow system. Pocket gophers create large numbers of fresh
mounds during certain peak periods of the year (Hickman & Brown 1973). Beetles may
orient to these mounds but the exact stimuli are unknown. The stimuli may be visual
(mounds contrast in color against the surrounding substrate), thermal (mounds absorb
and radiate heat at differing rates than the surrounding ground [unpublished data]),
chemical (rodent scent), or tactile (texture of old vs. fresh mounds). Beetles may find
a fresh active (with rodent) burrow which may be occupied by no other species or by
any of at least 6 other Aphodius species and any number of other commensal insects.
Many cave arthropods and burrow inhabitants have the wings and eyes absent or
poorly developed, and often have reduced pigment (e.g. the blind camel-cricket,
Typhloceuthophilus floridanus Hubbell 1940). None of the species treated here (with
the possible exception of A. aegrotus which does have reduced pigment) has evolved
these modifications, but other modifications have evolved. An elongation of appendages,
common in cavernicolous insects, is evident in their long narrow tarsi (most developed
December, 1991
Skelley & Woodruff: Five New Species of Aphodius (Coleoptera) 535
in A. tanytarsus). An expanded or explanate pronotum is present in several different
groups of Aphodius. Many of these Aphodius appear to be associated with burrowing
rodents.
The pocket gopher burrow and its associated fauna offer interesting opportunities
to study zoogeography, ecology, and speciation.
ACKNOWLEDGMENTS
We thank P. M. Choate, University of Florida, Gainesville, FL; P. W. Kovarik, Ohio
State University, Columbus, OH; R. W. Lundgren, Archer, FL; F. W. Skillman, Citra,
FL; M. C. Thomas, Florida State Collection of Arthropods, Gainesville, FL; and R. H.
Turnbow, Fort Rucker, AL, for their assistance in collecting these beetles, loans of
specimens, sharing of ideas, and reviews of this paper. We also thank Dr. G. Erdos and
D. Williams, Botany Department, University of Florida, for assistance with the SEM;
and Jeff Lotz, Florida Department of Agriculture, Division of Plant Industry, Gainesville,
FL, for photographic dark room work. We especially thank R. D. Gordon, United States
National Museum of Natural History, Washington, DC, for loans of related species,
review of the manuscript, and for his opinions and comments on these species. Lastly,
we thank the anonymous reviewers for their valuable comments. Florida Agriculture
Experiment Station, Journal Series No. R-01144.
REFERENCES CITED
BLUME, R. R., AND A. AGA. 1975. Aphodius from burrows of a pocket gopher in
Brazos County, Texas (Coleoptera: Scarabaeidae). Coleopt. Bull. 29(3): 161-162.
BLUME, R. R., AND A. AGA. 1979. Additional records of Aphodius from pocket gopher
burrows in Texas (Coleoptera: Scarabaeidae). Coleopt. Bull. 33(1): 131-132.
BLUME, R. R., AND J. W. SUMMERLIN. 1988. Histeridae (Coleoptera) from pocket
gropher burrows in east central Texas. Coleopt. Bull. 42(2): 202-204.
BROWN, W. J. 1928. The subgenus Platyderides in North America (Coleoptera). Canad.
Entomol. 60(1): 10-21; (2): 35-40.
CARTWRIGHT, O. L. 1939. Eleven new American Coleoptera (Scarabaeidae, Cicin-
delidae). Ann. Entomol. Soc. Am. 32: 353-364.
CARTWRIGHT, O. L. 1944. New Aphodius from Texas gopher burrows. Entomol.
News 55(5-6): 129-135, 146-150.
GORDON, R. D. 1983. Studies on the genus Aphodius of the United States and Canada
(Coleoptera: Scarabaeidae). VII. Food and habitat; distribution; key to Eastern
species. Proc. Entomol. Soc. Washington 85(4): 633-652.
HALDEMANN, S. S. 1848. Descriptions of North American Coleoptera, chiefly in the
cabinet ofJ. L. LeConte, M.D., with references to described species. Proc. Acad.
Nat. Sci. Philadelphia 1: 95-100.
HELAVA, J. 1978. A revision of the Neartic species of the genus Onthophilus Leach
(Coleoptera: Histeridae). Contrib. Am. Entomol. Inst. 15(5): 1-43.
HICKMAN, G. C., AND L. N. BROWN. 1973. Pattern and rate of mound production in
the Southeastern pocket gopher (Geomys pinetus). J. Mammol. 54(4): 971-975.
HORN, G. H. 1870a. Descriptions of the species of Aphodius and Dialytes of the United
States. Trans. Am. Entomol. Soc. 3: 110-134; pl. 3, legend p. 324.
HORN, G. H. 1870b. Descriptions of new species of Histeridae for the United States.
Trans. Am. Entomol. Soc. 3: 134-142, pl.I.
HORN, G. H. 1887. A monograph of the Aphodiini inhabiting the United States. Trans.
Am. Entomol. Soc. 14: 1-110.
HUBBELL, T. H. 1940. A blind cricket-locust (Typhloceuthophilus floridanus n. gen.
et sp.) inhabiting Geomys burrows in peninsular Florida (Orthoptera, Gryllac-
rididae, Rhaphidophorinae). Ann. Entomol. Soc. Am. 33: 10-32.
536 Florida Entomologist 74(4) December, 1991
HUBBELL, T. H. 1954. Relationships and distribution of Mycotrupes. In Olson, Hub-
bell, and Howden. The burrowing beetles of the genus Mycotrupes. Misc. Publ.
Mus. Zool. Univ. Michigan 84: 39-51.
HUBBELL, T. H. 1961. Endemism and speciation in relation to Pleistocene changes
in Florida and the southeastern coastal plain. Eleventh Int. Congress Ent., Wien
1960. 1: 466-469.
HUBBELL, T. H., AND C. C. GOFF. 1939. Florida pocket-gopher burrows and their
arthropod inhabitants. Proc. Florida Acad. Sci. 4: 127-166.
HUBBELL, T. H., A. M. LAESSLE, AND J. C. DICKINSON. 1956. The Flint-Chat-
tahoochee-Apalachicola region and its environments. Bull. Florida State Mus.
1(1): 1-72.
LECONTE, J. L. 1878. The Coleoptera of the alpine regions of the Rocky Mountains.
Art. XX. Bull. U.S. Geological and Geographical Survey of the Territories IV:
447-480.
ROBINSON, M. 1938. Studies in the Scarabaeidae. II. (Coleoptera). Trans. Am. En-
tomol. Soc. 64: 107-115.
Ross, E. S. 1940. New Histeridae (Coleoptera) from the burrows of the Florida pocket
gopher. Ann. Entomol. Soc. Am. 33(1): 1-9.
Ross, E. S. 1944. Onthophilus kirni new species, and two other noteworthy Histeridae
from burrows of the Texas pocket-gopher. Entomol. News 55: 115-118.
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the Rocky Mountains performed by order of Mr. Calhoun, Secretary of War,
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WOODRUFF, R. E. 1973. The scarab beetles of Florida. (Coleoptera: Scarabaeidae).
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1-220; 407 fig.
WOODRUFF, R. E. 1982. Rare and endangered Scarabaeidae of Florida, p. 84-102 in
R. Franz [ed.], Rare and endangered biota of Florida. Invertebrates. Vol. 6.
University of Florida Press, Gainesville, 131p.
SURVEY OF INSECTS OF SOUTH FLORIDA AND THE
FLORIDA KEYS: FLAT BARK BEETLES (COLEOPTERA:
CUCUJIDAE (SENS. LAT.) [LAEMOPHLOEIDAE:
PASSANDRIDAE: SILVANIDAE])
M. C. THOMAS
Florida State Collection of Arthropods
P. O. Box 147100
Gainesville, Florida 32614-7100 U.S.A.
STEWART B. PECK
Department of Biology, Carleton University
Ottawa, Ontario K1S 5B6 Canada
ABSTRACT
A survey of the flat bark beetles of tropical Florida (Dade, Monroe, Broward, and
Collier counties) revealed the presence of 26 species in three families: Laemophloeidae,
14 species; Passandridae, 2 species; Silvanidae, 10 species. Five species are newly re-
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Thomas & Peck: Flat Bark Beetles 537
corded for Florida and the United States: Laemophloeus suturalis Reitter,
Laemophloeus (sens. lat.) permixtus Grouvelle, Laemophloeus (sens. lat.) quinquear-
ticulatus Grouvelle, Monanus concinnulus (Walker), Nausibius sahlbergi Grouvelle,
Silvanus lewisi Reitter. One species, Rhabdophloeus horni (Casey), is newly recorded
from Florida. Collection localities and data are listed, and faunal affinities are discussed.
RESUME
Se llevo a cabo un muestreo de los escarabajos plans de la corteza en la region
tropical de Florida (condados de Dade, Monroe y Collier). Este muestreo indico la
presencia de 26 species incluidas en tres families: Laemphloeidae, con 14 species;
Passandridae, con dos species; Silvanidae, con 10 species. Se registran cinco species
por vez primera en Florida y en los Estados Unidos: Laemphloeus suturalis Reitter,
Laemophloeus (sens. lat.) permixtus Grouvelle, Laemophloeus (sens. lat.) quinquear-
ticulatus Grouvelle, Monanus concinnulus (Walker), Nausibius sahlbergi Grouvelle,
Silvanus lewisi Reitter. Se registra la especie Rhabdophloeus horni (Casey) por vez
primera en Florida. Se mencionan los datos y localidades de coleccion y se discuten las
afinidades de la fauna.
This paper is one of a series on the insect fauna of South Florida (Peck 1989). Data
were compiled from specimens collected by the second author at the approximately 30
sites detailed by Peck (1989), and from specimens examined by the first author in the
Florida State Collection of Arthropods (FSCA) and other collections. Additional sites
include: Camp Mahachee, a Girl Scout Camp adjacent to Matheson Hammock Park and
part of the same hammock; Ross Castellow Hammock, a Dade County park near Home-
stead; Upper Key Largo, that part of the key north of the intersection of U.S. 1 and
State Road 505 (much of this area is now part of a State Botanical Site and is protected
from further development). A total of 600 specimens-122 collected during Peck's survey
and 478 in the FSCA-forms the basis of this report. Peck's specimens were all collected
in large-area flight intercept traps unless otherwise indicated; FSCA specimens were
collected mostly in ultraviolet light traps or by hand.
There are many pending nomenclatural changes and unreported synonymies in these
families. However, currently accepted names are used here pending completion of a
faunal study for the entire state by the first author. Although it is generally agreed
that the traditional "Cucujidae" is a composite taxon of several genus groups that are
not particularly closely related to each other, the details of its division have not been
generally accepted. If the Cucujidae (sens. str.) is restricted only to Cucujus Fabricius
and related genera, then the Laemophloeinae must be raised to family rank, and it is
so treated here. Under that definition, no member of the family Cucujidae (sens. str.)
occurs in Florida.
RESULTS
Of the ninety-six described species of flat bark beetles occurring in America north
of Mexico (Thomas, unpublished), 56 occur in Florida. Of those, 26 are here recorded
from South Florida (Fig. 1). With the exception ofAhasverus advena (Waltl) and Cathar-
tus quadricollis (Guerin), no stored products pests were collected in the field during
this study and are not included in the total for South Florida, although they are included
in the totals for the state and for America north of Mexico. Three other species (Monanus
concinnulus (Walker), Cryptamorpha desjardinsi (GuBrin), and Silvanoprus scuticollis
(Walker)) are circumtropical in distribution and are undoubtedly introduced. Five species
Florida Entomologist 74(4)
East SE CA Intro./
Species US US NFla SFla Keys W.I. SA Cosmo.
Laemophloeidae
Dysmerus basalis X X X X
Laemophloeusfasciatus X X X X
Laemophloeusfloridanus X X X
Laemophloeus lecontei X X ?
Laemophloeus suturalis X X
Laemophloeus (sens. lat.)
bituberculatus X X X
Laemophloeus (sens. lat.)
permixtus X X X X
Laemophloeus (sens. lat.)
quinquerticulatus X X X
Lathropus pictus X X X
Lathropus vernalis X X X X
Placonotus macrognathus X X
Placonotus modestus X X X X X X
Placonotus politissimus X X X X X ?
Rhabdophloeus horni X
Passandridae
Catogenus rufus X X X X X X
Taphroscelidia linearis X X X X X
Silvanidae
Ahasverus advena X X X X X X X X
Ahasverus rectus X X X X
Cathartosilvanus trivialis X X X ?
Catharius quadricollis X X X X X X
Cryptamorphadesjardinsi ? X X X X
Monanus concinnulus X X X X
Nausibius sahlbergi X X ?
Silvanoprus scuticollis X X X X X X
Silvanus lewisi X X
Silvanus muticus X X X X
Fig. 1. Distribution of flat bark beetles recorded from South Florida and the Florida
Keys.
are newly recorded from Florida and the U.S., and one species is new to Florida.
Specimens are in the collections of the Florida State Collection of Arthropods, Canadian
Museum of Nature, and S. B. Peck.
Laemophloeidae
Dysmerus basalis Casey
This rarely collected species ranges from the Keys north to Alabama and the District
of Columbia. Ten specimens were seen from South Florida. Monroe Co.: Big Pine Key,
538
December, 1991
Thomas & Peck: Flat Bark Beetles 539
Key Largo, No Name Key, in May, June, and August, in ultraviolet light traps, flight
intercept traps, and reared from poisonwood, Metopium toxiferum (L.) Klug & Urban.
Laemophloeus (sens. lat.) bituberculatus Reitter
This distinctive species was first recorded from Florida by Thomas (1979). It ranges
from the Lower Keys (Stock Island) north to Highlands County, Lake Placid. It was
described from Puerto Rico and specimens also have been seen from the Bahamas.
Twelve South Florida specimens were seen as follows: Collier Co.: US41, 1.5mi. S. State
Road 951; Monroe Co.: Plantation Key, Stock Island, in January and April-May, in
ultraviolet light traps and under the bark of burned Myrica cerifera L.
Laemophloeus (sens. lat) permixtus Grouvelle
This species was described from Guadeloupe and occurs also in Cuba and the U.S.
Virgin Islands (St. Croix and St. Thomas). It is here newly recorded from Florida,
where it has been collected from the Keys north to Putnam County. Fifty-seven speci-
mens were seen from South Florida. Collier Co: Collier-Seminole State Park; Dade Co.:
Biscayne Bay, Camp Mahachee, South Miami (Old Cutler Hammock); Monroe Co.: Big
Pine Key (Cactus Hammock, Watson's Hammock), Big Torch Key, Cudjoe Key, Fat
Deer Key, No Name Key, Stock Is., Sugarloaf Key (Kitching Hammock), Key Vaca
(Marathon), from February to December, in flight intercept traps and in ultraviolet
light traps.
Laemophloeus (sens. lat.) quinquearticulatus Grouvelle
This distinctive species has been known previously only from southern Brazil. In
Florida it has been collected from Dade County north to Marion County. Only two
specimens from South Florida were seen. Dade Co.: Deering Estate Park, in May and
August, in ultraviolet light trap and evening car net.
Laemophloeus fasciatus Melsheimer
This primarily northeastern U.S. species ranges north to Canada. There is only one
South Florida record. Dade Co.: Miami, 2-IV-1961, in ultraviolet light trap.
Laemophloeus floridanus Casey
This species ranges north to North Carolina. One hundred forty-eight specimens
were seen from 21 collections in South Florida, but none has been seen from the Keys.
Dade Co.: Camp Mahachee, Castellow Hammock, Homestead, Miami, April to Sep-
tember, and December, in ultraviolet light traps.
Laemophloeus lecontei Grouvelle
This species is restricted presently to South Florida, although Laemophloeus chev-
rolati Grouvelle, described from Cuba, is probably conspecific. Twenty-nine South
Florida specimens were seen. Dade Co.: Biscayne Bay, Camp Mahachee, Matheson
Hammock; Monroe Co.: Upper Key Largo, from March to July, in ultraviolet light traps
and under the bark of gumbo limbo, (Bursera simarubra (L.) Sarg.).
Laemophloeus suturalis Reitter
This species is widespread in the Neotropics from Mexico south to Bolivia. We have
seen no specimens from the Antilles. Dade Co.: Castellow Hammock, Goulds, in April,
on dead, standing gumbo limbo (Bursera simarubra (L.) Sarg.).
Lathropus pictus Schwarz
This maculate species is found only in southern Florida and the Antilles. Forty-one
specimens were seen. Dade Co.: Camp Mahachee, Deering Estate Park, from April to
June, and August to September, in a flight intercept trap and in ultraviolet light traps.
Lathropus vernalis Casey
All of the non-maculate, dark specimens of Lathropus are included under this name,
although there is probably more than one species represented. It ranges north to Canada
Florida Entomologist 74(4)
and west to Oklahoma. Twelve specimens from South Florida were seen. Dade Co.:
Camp Mahachee; Monroe Co.: Plantation Key, Upper Key Largo, from April to May,
and September, in ultraviolet light traps.
Placonotus macrognathus Thomas
This species was described from Upper Key Largo and is known only from the type
series and a specimen from Cuba (Thomas 1984). Monroe Co.: Upper Key Largo, April
and May, in ultraviolet light trap and under the bark of gumbo limbo, (Bursera simarubra
(L.) Sarg.).
Placonotus modestus (Say)
This widespread species ranges from New York west to Arizona and south to Panama,
as well as Cuba, Jamaica, and Trinidad. It has not been collected on the Keys. Twenty-one
South Florida specimens were seen. Dade Co.: Biscayne Bay, Camp Mahachee, Castellow
Hammock, Everglades National Park (Long Pine Key), Goulds, South Miami (USDA
Subtrop. Res. Sta.), from April to June, and August, in flight intercept traps, ultraviolet
light traps, and under the bark of Australian pine, Casuarina sp.
Placonotus politissimus (Wollaston)
This species occurs both in the Afrotropical region and the Neotropics. In Florida
it ranges from the Keys north to Marion County. Twenty-seven specimens were seen
from South Florida. Dade Co.: Camp Mahachee, Everglades National Park (Long Pine
Key), Goulds, Homestead, Matheson Hammock, Miami (USDA Plant Intro. Gardens);
Monroe Co.: Stock Island, Upper Key Largo, from January to February, April to June,
August to September, and December, in flight intercept traps and in ultraviolet light
traps.
Rhabdophloeus horni (Casey)
This species, known previously from California and Arizona, is here newly recorded
from Florida, where it occurs only in the Upper Keys. Five specimens from South
Florida were seen. Monroe Co.: Upper Key Largo, from April to June, in ultraviolet
light traps and under the bark of gumbo limbo, (Bursera simarubra (L.) Sarg.).
Passandridae
Catogenus rufus (Fabricius)
This widespread species ranges north Canada and west to Texas. It probably occurs
throughout Florida. Sixty-six specimens were seen from South Florida. Collier Co.: no
locality; Dade Co.: 2 mi. SW of Hallandale, Coral Gables, Everglades National Park
(1.5km W Royal Palm and Long Pine Key), Perrine, Richmond; Monroe Co.: Big Pine
Key (Watson's Hammock), Key Largo (Pennekamp State Park and Upper Key Largo),
Little Torch Key, Plantation Key, Stock Island, from January to August, and November,
in flight intercept traps, ultraviolet light traps, and under the bark of dead Quercus,
Pinus, and Lysiloma.
Taphroscelidia linearis (LeConte)
This species has been reported from Florida and Texas south to Brazil. The species
of this genus are difficult to distinguish and the records may be unreliable. In Florida
it probably occurs throughout the state. No specimens from the Keys were seen during
this study, but Schwarz (1890) recorded it from Key West. Only one specimen was seen
from South Florida. Dade Co.: Perrine, 5-6-V-1977, ultraviolet light trap.
540
December, 1991
Thomas & Peck: Flat Bark Beetles
Silvanidae
Ahasverus advena (Waltl)
This is a cosmopolitan pest species of stored products that is sometimes collected in
the wild. Five specimens were seen from South Florida. Dade Co.: Miami; Monroe Co.:
Stock Is., in April and June, in a lumber warehouse and under debris.
Ahasverus rectus (LeConte)
This commonly collected, widespread species occurs from Florida north to North
Carolina and west to Arizona. One hundred twenty-three specimens were seen from
South Florida. Collier Co.: Collier-Seminole State Park; Dade Co.: Camp Mahachee,
Castellow Hammock, Everglades National Park (Long Pine Key), Florida City, Mathe-
son Hammock, Miami (USDA Subtropical Res. Sta.); Monroe Co.: Bahia Honda State
Park, Dry Tortugas (Garden Key), Fat Deer Key, Key Largo (Pennekemp State Park
and Upper Key Largo), Stock Island, Sugarloaf Key, Key Vaca, from March to Sep-
tember, and November, in flight intercept traps, ultraviolet light traps, and by sampling
forest floor litter in a Berlese funnel.
Cathartosilvanus trivialis (Grouvelle)
This widespread Neotropical species was first recorded from Florida by Thomas
(1979). It occurs throughout the Neotropics, ranging north in California and Arizona in
the West and into South Florida in the East. Six specimens were seen from South
Florida. Broward Co.: Wilton Manors; Dade Co.: Goulds, Miami Springs, in February,
April and June, under bark of Casuarina sp.
Cathartus quadricollis (Gu6rin)
This species is a pest especially of corn, both stored and in the field. It is nearly
cosmopolitan in distribution and occurs in the U.S. from Florida west to California and
north to the District of Columbia. Eleven specimens were seen from South Florida.
Dade Co.: Homestead, Naranja, in April and August, in ultraviolet light traps, on
Cattleya sp., and in decaying seed pods of tamarind, Tamarindus indica L.
Monanus concinnulus (Walker)
This species is circumtropical in distribution. It is here reported from the U.S. for
the first time based on two South Florida specimens. It is uncertain whether this species
is established in Florida. Dade Co.: Coral Gables, in March, on Bambusa vulgaris
Schrad. ex J. C. Wendl.
Nausibius sahlbergi Grouvelle
This species was described from Brazil and is here recorded from Florida for the
first time, based on two specimens in the FSCA from Dade County collected by H. F.
Strohecker in May, 1949 and two specimens in the American Museum of Natural History
collected by A. M. Nadler at Royal Palm Park on 14 February 1957.
Silvanoprus scuticollis (Walker)
This circumtropical species was first recorded from the U.S. by Thomas (1979). It
occurs in the southeastern U.S. north to West Virginia and west to Oklahoma. Twenty-
seven specimens were seen from South Florida. Dade Co.: Camp Mahachee, Matheson
Hammock, Perrine, Homestead, in January, April to July, and September, in ultraviolet
light traps.
Silvanus lewisi Reitter
This species is widespread in the Old World tropics and is here recorded from Florida
and the U.S. for the first time, based on two specimens in the FSCA from South Florida.
Florida Entomologist 74(4)
Broward Co.: Davie; Dade Co.; no locality, in February, in a building. It is uncertain
whether this species is established.
Silvanus muticus Sharp
This is a widespread eastern U.S. species (Halstead 1973). It has not been collected
in the Keys. Only one specimen was seen from South Florida. Dade Co.: Camp Mahachee,
27-V-1983, ultraviolet light trap.
Cryptamorpha desjardinsi (Gubrin)
This species is circumtropical in distribution. In the U.S. it also has been reported
from Alabama (Loding 1945). In Florida it is restricted to the southern half of the state,
ranging north to Orange County. Seventeen specimens were seen from South Florida.
Dade Co.: Camp Mahachee, Coral Gables, Goulds, Miami, from March to July. Specimens
have been collected in ultraviolet light traps, and on Saccharum officinarum L., Bam-
busa vulgaris Schrad. ex J. C. Wendl, and "orchids".
BIOGEOGRAPHICAL CONSIDERATIONS
Fig. 1 presents a summary of the distributions of the species known to occur in South
Florida.
Of the 26 species here recorded from South Florida, six species (Ahasverus advena,
Silvanus lewisi, Silvanoprus scuticollis, Monanus concinnulus, Cathartus quadricollis,
and Cryptamorpha desjardinsi) are clearly introduced. Curiously, all are members of
the Silvanidae. There are no precinctive species of these three families in Florida.
Of the remaining silvanids, two (Cathartosilvanus trivialis and Nausibius sahlbergi)
are Neotropical and probably introduced; one (Ahasverus rectus) is widely distributed
in the southern U.S. and probably invaded South Florida from the North, although its
affinities are Neotropical; and one (Silvanus muticus) is Nearctic in distribution and
affinities and probably invaded from the North.
It is only among the Laemophloeidae that an Antillean faunal element manifests
itself. Laemophloeus (sens. lat.) bituberculatus, Laemophloeus lecontei, Laemophloeus
(sens. lat.) permixtus, Lathropus pictus, and Placonotus macrognathus are clearly
Antillean in origin and affinities. Of the other species, Placonotus politissimus is prob-
ably introduced from Africa and Laemophloeusfasciatus is Nearctic in distribution and
probably invaded from the North; the remainder are either widespread in the Neotropics
and probably invaded South Florida by way of the northern coast of the Gulf of Mexico,
or are widespread in both the Neotropics and southeastern U.S. and invaded South
Florida from the North.
ACKNOWLEDGMENTS
Field work of Stewart Peck was partly supported by an operating grant from the
Natural Sciences and Engineering Council of Canada. We would like to thank Carl Ostl
of the Girl Scout Council of Tropical South Florida for permission to collect at Camp
Mahachee and Lori Parker and John Gleason for tending a light trap there. We also
thank G. B. Edwards, Avas B. Hamon, Willis W. Wirth, R. E. Woodruff, and an
anonymous reviewer for their constructive criticisms of the manuscript, and Carlos R.
Artaud for translating the abstract. This is Contribution No. 759, Bureau of Entomology,
Division of Plant Industry, Florida Department of Agriculture & Consumer Services.
REFERENCES CITED
HALSTEAD, D.G.H. 1973. A revision of the genus Silvanus Latreille (s.l.) (Coleoptera:
Silvanidae). Bull. British Mus. Natur. Hist. (Entomol.) 29: 39-112.
December, 1991
Roach: Overwintering sites 543
LODING, H. P. 1945. Catalogue of the beetles of Alabama. Geol. Surv. Alabama,
Monograph 11: 1-172.
PECK, S. B. 1989. A survey of insects of the Florida Keys: post-Pleistocene land-bridge
islands: Introduction. Florida Entomol. 72: 603-612.
SCHWARZ, E. A. 1890. [Coleoptera of Key West]. Proc. Ent. Soc. Washington 1: 93-94.
THOMAS, M. C. 1979. Flat bark beetles new to Florida and the U.S. (Coleoptera:
Cucujidae s.l.). Coleops. Bull. 33: 357-358.
THOMAS, M. C. 1984. A revision of the New World species of Placonotus Macleay
(Coleoptera: Cucujidae: Laemophloeinae). Occ. Pap. Florida St. Coll. Arthr. 3:
vii + 28pp.
NATURAL PLANT MATERIALS AS OVERWINTERING SITES
FOR ARTHROPODS IN THE COASTAL PLAIN OF
SOUTH CAROLINA
S. H. ROACH
USDA-ARS, Cotton Production Research Unit
P. 0. Box 2131, Florence, SC 29503
In cooperation with the South Carolina Agricultural
Experiment Station
ABSTRACT
Six types of plant materials, grass hay, wheat straw, pine needles, woods litter, corn
stalks and husks, and broomsedge grass clumps were compared in small plots as sites
for overwintering arthropods. Significant differences were found between numbers of
some insects in the materials, with the coccinellids primarily overwintering in the broom-
sedge clumps, while Staphylinids preferred Bermuda grass hay and wheat straw. Overall,
a limited number of important predaceous and phytophagous species were found in the
different materials. There were significant differences in the number of arthropods
depending upon where the materials were placed in relation to open field or field edges.
RESUME
Se compararon en parcelas pequefias seis tipos de materials de plants, heno de
pastos, paja de trigo, agujas de pino, rastrojo de arboles, cafia y chalas de maiz, y past
escoba de junco, como lugares de hibernacion de artropodos. Se encontraron diferencias
significativas entire los materials en los numerous de algunos insects, con los coccinelidos
hibernando primeramente en el past escoba de junco, mientras que los Staphylinidos
prefirieron el heno de past Bermuda y la paja de trigo. Sobretodo se encontro, un
numero limitado de predatores importantes y species fitofagos en los diferentes
materials, dependiendo del sitio donde se colocaron los materials en relacion con un
campo abierto on con el margen del campo.
In temperate and cold climates, arthropods have evolved a number of strategies for
surviving the winter months. The habitats chosen by arthropods for overwintering are
quite variable but relatively specific for individual species. Some of the early work on
arthropod overwintering in the United States was done by Blatchley (1895) in Indiana
Florida Entomologist 74(4)
and Holmquist (1926) in Illinois. They made detailed observations of individual species
and their location during the winter months. Rainwater (1941) listed several hundred
species of insects that were collected from samples of woods trash, Spanish moss, cotton
gin trash, and wild cotton. The majority of specimens were found in the woods trash
samples. Kirk & Taft (1970) identified 405 species of Coleoptera collected during the
winter in South Carolina from woods trash samples. These included 35 well-known crop
pests and over 100 species from families that are mostly predaceous. Jones and Sullivan
(1981) determined the overwintering sites and spring emergence patterns of 47 hemip-
terous species found in woods trash and open field habitats in South Carolina.
In Florida, Plagens & Whitcomb (1986) examined corn residues for overwintering
arthropods and found 24 species of spiders and 25 species of predaceous Coleoptera,
Hemiptera and Dermaptera. Magnolia leaf litter was found to provide overwintering
sites for a large number of arthropods in Pennsylvania, including 25 heteropterans such
as Lygus lineolaris (Palisot de Beauvois) and Nabis roseipennis Reuture (Wheeler &
Stimmel 1988). The overwintering behavior of the predaceous Coccinellidae has been
reviewed by Hagen (1962) and Hodek (1967).
There are numerous other articles in the literature on the overwintering habits of
specific arthropods, but few that deal with habitat conservation and man-made or pro-
vided habitats for overwintering arthropods. In England, a great deal of research has
been conducted on the relationship between winter crops, uncultivated areas, and
hedgerows on the occurrence and distribution of pest and predatory arthropods (van
Emden 1963, 1965, Lewis 1965, Sotherton 1984). Their efforts have shown direct relation-
ships between habitats and ensuing populations of arthropods.
In the United States, Tamaki & Halfhill (1968) used bands of sheet aluminum lined
with heavy paper and tarred burlap around the main branches of peach trees to provide
shelter for predators of the green peach aphid. They found that several species of
predators successfully used the bands for overwintering, but only a few pest insects
used the bands for shelter. In a similar experiment, Mizell & Schiffhauer (1987) tested
burlap bands, Coolpad* filters, layered cardboard and natural tree bark as overwintering
sites for predators in pecan orchards. They reported all of the habitat types were useful
for monitoring some arthropods but none appeared useful for augmenting overwintering
populations. In another study, Fye (1985) constructed traps of layered corrugated fiber-
board that effectively substituted for natural overwintering sites for some predators in
pear orchards.
The purpose of the present study was to determine if several native materials located
in field crop areas in South Carolina could serve as overwintering sites for arthropods
and to record the important species of predaceous and phytophagous arthropods that
were present in each type of material.
MATERIALS AND METHODS
A 6.9-ha field surrounded by largely undisturbed mixed deciduous and pine woodlands
on the Pee Dee Research and Education Center, Florence, South Carolina was chosen
for this study. The field is primarily used for experimental plots of cotton and tobacco
with areas of mixed weeds and grasses. In September 1987, 4 replicates of 6 different
plant materials-wheat straw, mixed deciduous woods litter, pine straw, Bermuda grass
hay, corn stalks and husks, and broomsedge plants (Andropogen virginicus L.) (8 plants/
plot-were installed in the middle (2 replicates) and along the SW side (2 replicates) of
the field. Each plot of plant materials within a replicate was assigned position at random
and separated by 4.5 m. Replicates were approximately 100 m apart in all directions.
Plant materials were piled to an approximate depth of 15 cm, except for the broom-
sedge plants, which were 15-20 cm in base diameter and approximately 0.75 m tall when
December, 1991
Roach: Overwintering sites
545
transplanted into the plots. No retaining barriers were used around the plots, but
frequent checks were made during the winter to ensure that the plots were not scattered
by animals or weather. The experiment was repeated in the same field in 1989-90, but
with 2 more replications of each plot added so that there were 3 plots of each of the 6
types of plant material in the middle of the field, and 3 plots along the SW edge of the
field. During both years the field was left fallow following crop harvest and removal of
the plot materials for examination. Plots were established between 22 June and 20
September, whenever the various materials were available or best suited for transplant-
ing (broomsedge).
During the period 27 January-8 March 1988 and 12 December 1989-1 March 1990,
the plant materials within each plot were collected down to bare soil, placed in cloth
bags, and brought into a field laboratory for examination. The plant materials were
examined on a steel-topped table underlayed with electric resistance heat coils that
maintained a temperature of approximately 44C. Examination time varied tremendously
between the different materials due to the materials themselves and moisture content
when collected, but normally samples were held on the table until the material temper-
ature reached that of the table, and/or no further arthropods were found in the material.
All arthropods emerging or found alive in the materials were placed in 70% ethyl alcohol
until they could be identified. Plot materials were kept in the bags in a large walk-in
cooler maintained at 7+ 2C until they could be examined. The arthropod collections
were sorted and identified to groups or common species, and the number occurring in
the various materials were compared using Fisher's Protected LSD at P=0.05 after
data transformation using the formula n +0.5. Data transformation was performed to
allow analysis between groups of arthropods that varied widely in numbers in the
different overwintering materials and locations. Major arthropod groups such as the
spiders, carabids, etc. were grouped for analysis and most species were not identified
to species unless indicated. Specimens identified to species are deposited in the collection
of Clemson University at the Pee Dee Research and Education Center, Florence, South
Carolina. Unless otherwise noted, all identifications were performed by the author.
RESULTS AND DISCUSSION
A great many arthropod species were found overwintering in the different plant
materials, and significant differences occurred between overall numbers of arthropods,
individual insect groups, species, and plot location. Since the purpose of the experiment
was to record the overwintering of predaceous and phytophagous arthropods of impor-
tance in field crops, many species observed were omitted from the results.
1987-88 Study
Interaction between the location and plant materials within the field affected the
numbers of arthropods, especially the coccinellids, Hippodamia convergens (Guerin-
Meneville) and Coccinella septempunctata (L), that overwintered in some plots. Broom-
sedge located in full-day sunlight held the largest number of overwintering H. con-
vergens, and total arthropods collected (due to the higher number of H. convergens
found) (Table 1). Plots located adjacent to the woods that received a half day or less of
sunlight consistently held less coccinellids than the fully exposed plots. Only a few H.
convergens were found in the other five types of plant materials, regardless of where
they were located. Coccinella septempuncata is an introduced species that became
established in the South Carolina Coastal Plain area in 1984 (personal observations).
This species also preferred broomsedge and was not found in any other materials except
pine straw.
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Other predaceous arthropods found in the plots are shown in Table 2. Coleomegilla
maculata (De Geer) occurred most frequently in broomsedge and only a few were found
in any of the other materials. The carabids were found in broomsedge and pine straw
more than the other materials except for woods litter. In previous surveys of undisturbed
woods litter Kirk & Taft (1970), and Rainwater (1941) reported 47 and 53 species of
carabids, respectively, as occurring in the Coastal Plain area of South Carolina. The
Staphylinids occurred significantly more often in Bermuda grass hay. This is another
large family with 50 species previously reported by Kirk & Taft (1970) and 37 species
by Rainwater (1941) from the Coastal Plain area. A few Scymnus beetles, primarily S.
loewii and S. creperus were found overwintering in 3 of the plots, but numbers were
insignificant.
The spiders as a group were fairly evenly distributed among the plots, as were the
harvestmen (primarily Hadrobunus spp.), Geocoris, spp., and reduviids. Many of the
specimens of these groups collected except Geocoris, were immature, and species iden-
tifications were not attempted. A few phytophagous species of insects were found in
the plots, but none of the known crop nests were found in significant numbers in any
habitat (Table 3). This was somewhat unexpected since several species of pentatomids,
L. lineolaris, and many other phytophagous species are known to overwinter in woods
litter (Rainwater 1941). Only one rice stink bug, (Oebalus pugnax (Fabricius) and one
Thyanta custator (Fabricius) were taken from the plots (both from woods trash), and
no L. lineolaris were found in any of the materials. Crickets were found in all the
different plant materials but slightly more frequently in the pine straw, woods litter
and broomsedge.
Considering the total numbers of arthropods collected from the different plant mate-
rials, the broomsedge, primarily because of the large number of coccinellids, held the
highest number of overwintering arthropods, followed by the Bermuda grass hay. How-
ever, excluding broomsedge, the other 5 plant materials harbored similar numbers of
arthropods. Overall, the corn residues appeared to be the least attractive, since no
groups or species inhabited this residue in significantly greater numbers than the other
plant materials.
1989-90 Results
Interaction of location and plant materials occurred again in the 2nd year of the
study, but the number and species of insects involved were different (Table 4). Coccinella
septempunctata again showed distincly higher numbers in the unshaded broomsedge as
did Geocoris spp., L. lineolaris and the other miscellaneous Heteroptera. Hippodamia
convergens was present in much smaller numbers during 1989-90, but broomsedge con-
tained higher numbers than the other materials (Table 5). High variability between the
broomsedge locations is characteristic of this species since it aggregates in only a few
sedge clumps in a field area regardless of the number of apparently suitable sites avail-
able. Coleomegilla maculata was present in slightly higher numbers than in the previous
year of this study, and the broomsedge had significantly more beetles than the other
materials. The carabids as a group also showed similar distributions in the materials as
in 1987-88, with a fairly general scattering among the materials, Staphylinids were much
more generally distributed in 1989-90 than in 1987-88, and no clear site preferences were
noticeable. Scymnus spp. occurred in very low numbers during both years of the study
and were not numerous enough to compare differences in locations in 1989-90.
Spiders as a group were more numerous the second year of the study in all plots,
but little differences in number was noticeable except for a scarcity of spiders in the
corn residues. The same was true of the harvestmen, which was different from 1987-88
only in that broomsedge was also populated by similar numbers as the other materials.
Roach: Overwintering sites
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Roach: Overwintering sites
Reduviids were found in low numbers in all materials in 1989-90 while nabids were more
frequently collected than in the previous year. The nabid genera Reduviolus and Hop-
listoceles were more frequently collected in 1989-90 and were responsible for the increases
shown in Table 5.
Photophagous insects and total arthropods found in the various plant materials are
shown in Table 6. Low numbers of pest insects were present in all the plant materials,
just as in 1987-88. Most of the stink bug species collected are usually not serious crop
pests (Mormidea spp.; Thyanta spp.) and the miscellaneous hemipteran species such as
Pachybrachia and Ischnodemus are largely unknown as pests of agricultural crops. The
vegetable weevil (Listroderes costerostris obliquus Klug) is a serious pest of many field
crops and was relatively evenly distributed among the overwintering habitats and oc-
curred in fairly low numbers.
Total arthropod numbers found in the various plant material plots were also similar
to those found in 1987-88, except in magnitude. Overall, more arthropods were found
in the broomsedge than any material except pine straw, while corn stalks and leaf litter
remained low in numbers similar to the first year of the study.
The inability of the small plots of plant materials to attract many of the major
predaceous or phytophagous species could be due to a number of factors, such as proxim-
ity to the much more expansive woodlands that satisfy climatic preferences such as
shading, moisture level, and depth of the soil humus layers, all of which moderate
weather changes. Coccinellid species that aggregated, such as H. convergens and C.
maculata may exhibit hypsotaxis as well as chemotaxis which attracts them to more
specific microhabitats such as the unshaded broomsedge clumps or isolated trees with
thick layers of soil humus. In areas with limited woodlands or uncultivated areas, plots
or strips of various decomposing plant materials could be more attractive to a wider
range of arthropods. Some of the coccinellids, however, are apparently aided in overwin-
tering by the presence of thick broomsedge clumps in open areas in the coast plain of
South Carolina.
ACKNOWLEDGMENTS
Special appreciation is extended to Dr. Merrill Sweet for identifying the Geocoris
spp., to Dr. James Cokendolpher for identifying the Opiliones, and to Dr. R. D. Gordon
for identifying the Scymnus spp. In cooperation with the South Carolina Agricultural
Experiment Station, this paper reports research results only. Mention of a commercial
or proprietary produce does not imply endorsement by the USDA-ARS.
REFERENCES CITED
BLATCHLEY, W. S. 1895. Notes on the winter insect fauna of Vigo County, Indiana
I. Orthoptera. Psyche VII 248-250.
FYE, R. E. 1985. Corrugated fiberboard traps for predators overwintering in pear
orchards. J. Econ. Entomol. 78: 1511-1514.
HAGEN, K. S. 1962. Biology and ecology of predaceous Coccinellidae. Ann. Rev.
Entomol. 6: 289-326.
HODEK, I. 1967. Bionomics and ecology of predaceous Coccinellidae. Ann. Rev. En-
tomol. 12: 79-104.
HOLMQUIST, A. M. 1926. Studies in arthropod hibernation. I. Ecological survey of
hibernating species from forest environments of the Chicago region. Ann. En-
tomol. Soc. Amer. XIX. 395-428.
JONES, W. A., AND M. J. SULLIVAN. 1981. Overwintering habitats, spring emergence
patterns, and winter mortality of some South Carolina Hemiptera. Environ. En-
tomol. 10: 409-414.
Florida Entomologist 74(4)
KIRK, V. M., AND H. M. TAFT. 1970. Beetles found in woods trash during winter boll
weevil surveys. USDA, ARS Prod. Res. Rept. 199. 12 p.
LEWIS, T. 1965. The effects of shelter on the distribution of insect pests. Sci. Horticul-
ture. 17: 74-84.
MIZELL, R. F. III, AND D. E. SCHIFFHAUER. 1987. Trunk traps and overwintering
predators in pecan orchards: Survey of species and emergence times. Florida
Entomol. 70: 238-244.
PLAGENS, M. J., AND W. H. WHITCOMB. 1986. Corn residue as an overwintering
site for spiders and predaceous insects in Florida. Florida Entomol. 69: 665-671.
RAINWATER, C. F. 1941. Insects and spiders found in Spanish moss, gin trash, and
woods trash, and in wild cotton. USDA, Bureau of Entomol. and Plant Quarantine.
E-528. 20 p.
SOTHERTON, N. W. 1984. The distribution and abundance of predatory arthropods
overwintering on farmland. Ann. Appl. Biol. 104: 423-429.
TAMAKI, GEORGE, AND J. E. HALFHILL. 1968. Bands on peach trees as shelters for
predators of the green peach aphid. J. Econ. Entomol. 61: 707-711.
VAN EMDEN, H. F. 1963. A preliminary study of insect numbers in field and hedgerow.
Entomol. Monthly. 98: 255-259.
WHEELER, A. G., JR., AND J. F. STIMMEL. 1988. Heteroptera overwintering in
magnolia leaf litter in Pennsylvania. Entomol. News. 99: 65-71.
HOSTS OF A PHONOTACTIC PARASITOID AND
LEVELS OF PARASITISM
(DIPTERA: TACHINIDAE: ORMIA OCHRACEA)
T. J. WALKER AND S. A. WINERITER
Department of Entomology and Nematology
University of Florida
Gainesville, Florida 32611-0740
ABSTRACT
In central Texas, females of Ormia ochracea (Bigot) find hosts, Gryllus integer
Scudder (Orthoptera), by homing on the hosts' calling songs. 0. ochracea is abundant
each fall in Alachua County, Florida, where G. integer does not occur. Gryllus rubens
Scudder and G. firms Scudder were collected by three methods and held for emergence
of 0. ochracea. Of 185 G. rubens and 100 G. firms collected during January to August,
none were parasitized. Levels of parasitism during the remaining months never exceeded
10% except at sound-baited traps-but these attracted larvipositing 0. ochracea as well
as G. rubens. For specimens collected in pitfall traps and by searching under objects,
levels of parasitism of G. rubens and G. firmus and of males and females were unexpec-
tedly similar-though 0. ochracea is not attracted to firmus calls and female Gryllus
do not call. When muted and nonmuted reared males of G. rubens were experimentally
exposed in the field in the fall for 5 days, no muted males were parasitized but 7 of 13
nonmuted males were.
RESUME
En la parte central de texas, las hembras de Ormia ochracea (Bigot) encuentran sus
hospederos, Gryllus integer Scudder (Ortoptera) al ser atraidos por el huesped. 0.
ochraceae es abundante cada otofio en el condado Alachua, Florida, donde no se encuentra
December, 1991
Walker & Wineriter: Hosts of a Phonotactic Parasitoid 555
G. integer. Se colectaron Gryllus rubbens Scudder y G. firmis mediante el uso de 3
metodos y se mantuvieron para esperar la emergencia de 0. acraceae. Se colectaron
185 G. rubens y 100 G. firmus entire Enero y Agosto y ninguno fue parasitado. Los
niveles de parasitismo durante los meses restantes nunca excedieron mas del 10% except
en trampas con sonido como atrayente; sinembargo, estas atrayeron no solo 0. ochraceae
sino tambien G. rubens. Para los especimenes colectados en trampas de lata enterradas,
los niveles de parasitismo de G. rubens y G. firmus y de machos y hembras fueron
similares; aunque 0. ochraceae no es atraida hacia las llamadas defirmus y las hembras
de Gryllus no emiten sonidos de llamada. Cuando machos callados de G. rubens y machos
que emitian sonidos fueron expuestos expirementalmente por 5 dias en el campo durante
el otofio, los machos que emitian sonidos fueron parasitados y 7 de los 13 machos callados
tambien lo fueron.
At least three species of ormiine tachinids are phonotactic parasitoids of ensiferan
Orthoptera (Cade 1975, Burk 1982, Fowler & Kochalka 1985). Interest in the host ranges
of these flies and levels of parasitism is sharpened by the recent release in Florida of
Ormia depleta (Wiedemann), a South American species, for the control of pest mole
crickets (Scapteriscus spp.) (Wineriter & Walker 1990).
Ormia ochracea (Bigot) is one of six ormiines native to the southeastern United
States (Sabrosky 1953a, b). Cade (1975), working in Travis County, Texas, discovered
that its larvipositing females were attracted to the taped song of Gryllus integer Scudder.
Cade (1979) reported that 15 of 58 G. integer males he collected were parasitized (26%).
Of the 14 males collected by their calls, 11 (79%) were parasitized; of 44 males collected
by other means, 4 (9%) were parasitized; of 21 females (which do not call), none was
parasitized. Cade (1984) reported 17 of 73 G. integer calling males were parasitized
(23%), whereas none of the 17 or more males collected under lights were parasitized.
These numbers suggest that 0. ochracea females find few hosts except by their calls.
Mangold (1978), working in Alachua County, Florida, attracted small numbers of gravid
0. ochracea females to the taped song of Scapteriscus borellii Giglio-Tos (= S. acletus
Rehn & Hebard: Nickle 1992). He found that larvae dissected from an 0. ochracea
female could develop in individuals of S. borellii and Gryllus rubens Scudder, but noted
that parasitoids can often be propogated on unnatural hosts. Walker (1986, 1989, and
unpublished), in studies at one of Mangold's sites, attracted ca. 1,000 female 0. ochracea/
trap/yr to traps baited with the calling song of G. rubens. (G. rubens is the southeastern
field cricket most closely related to G. integer). More than 80% of the sound trap captures
were in September through December. Wineriter and Walker (1990) maintained a lab-
oratory colony of 0. ochracea on Gryllus rubens, and showed (1990 and unpublished)
that larvae could develop on Gryllus firmus Scudder, Gryllus ovisopis Walker, Acheta
domesticus (L.), Scapteriscus borellii, and last instar juveniles of G. rubens.
We started this study to determine how much 0. ochracea contributed to the mortality
of adult G. rubens. The abundance of 0. ochracea in the fall suggested high levels of
parasitism, as did Burk's (1982) data on Ormia lineifrons, a phonotactic parasitoid of
Neoconocephalus triops (L.) (40-90% parasitized, n>39). As a control to our phonotactic
biases, we included G. firmus, a field cricket common in the same habitats as G. rubens
but whose song does not attract 0. ochracea (Walker 1986).
METHODS AND MATERIALS
Our first method was to collect Gryllus spp. adults and hold them 10 days, a time
sufficient for 0. ochracea larvae to complete development, emerge, and pupate (Wineriter
& Walker 1990). Three collecting techniques were used: sound-baited traps in pastures
Florida Entomologist 74(4)
and wooded areas (Walker 1986), linear pitfall traps in pastures (Lawrence 1982), and
turning over objects and disturbing litter ("grubbing") in an organic garden. The sound
traps broadcast synthetic rubens and firmus songs and were designed to catch flying
crickets as they landed at the sound. About 40% of sound-trapped crickets are males
(Walker 1986).
Because parasitized crickets might not fly, thus biasing our sound trap results, we
determined parasitization of flying and non-flying crickets in the laboratory. At a sound
trap that was heavily visited by 0. ochracea we captured 360 G. rubens and placed
them in flight test cages that could be escaped only by flying (Walker 1987). We held,
for parasite emergence, 79 crickets that flew from the test cage during the next five
nights. We also held a matching sample of 79 crickets that had remained in the test cage.
Our second method for studying natural parasitism was to expose parasite-free,
caged G. rubens males for 5 days in an 0. ochracea infested pasture. The males were
reared in the laboratory and half were muted before exposure by cutting off their right
tegmen. Long-winged males were rendered flightless by cutting off one of their
matathoracic wings. The males were exposed individually in a circle of 12, evenly spaced
35 x 28 cm (h x dia) plastic buckets. Muted and nonmuted males were alternated. Each
bucket had 5 cm of moist sand and a plug of sod at the bottom to provide shelter for
the cricket, and a cover of 12-mm mesh hardware cloth to allow flies to enter while
excluding larger enemies. After 5 days the males were taken to the laboratory and held
for emergence of parasites; some of those that died during the 10-day holding period
were dissected to determine if Ormia larvae were present. Four replicates were run
between 22 Sep and 15 Oct 1988.
RESULTS AND DISCUSSION
Sound trapping. Sound-trapped G. rubens were heavily parasitized in the fall (Table
1). However, numerous gravid females of 0. ochracea are attracted to rubens sound
traps in the fall (Walker 1986, 1989), suggesting that crickets may have been parasitized
after they landed at the trap. Thus we checked the amount of time elapsed between
sound-trapping and emergence of parasite larvae from their hosts. Emergence within
6 days would indicate that the crickets were already parasitized when they arrived at
the sound trap (Wineriter and Walker 1990). For 61 rubens trapped Sep-Dec 1986, 1987,
our records of emergence were adequate to apply the 6-day criterion. No larvae had
emerged during the first 6 days-i.e., all hosts may have been parasitized after reaching
the sound trap.
These results showed the pertinence of our laboratory study about whether
parasitized crickets fly. Of the 79 crickets that flew from test cages, 67 (85%) proved
to be parasitized by 0. ochracea. Of 79 crickets that did not fly, 69 (87%) were parasitized.
We concluded that parasitized crickets flew and in proportions similar to non-parasitized
crickets. In fact, five parasitized crickets flew when the larvae they hosted were well
advanced: larvae emerged in 3-4 days, whereas total larval development generally takes
7-8 day.
More sound-trapped G. rubens were monitored in the fall of 1990, and about half
were parasitized by 0. ochracea. In 3 cases the parasite emerged in less than 7 days
(Table 1), suggesting a prior-to-trapping level of parasitism similar to that revealed by
other methods.
Pitfalls and grubbing. As in the case of sound traps, all parasitism was in the fall
(Table 1). Of 100 Gryllus captured in pitfalls in the fall, 2 rubens and 0 firmus were
parasitized by 0. ochracea. One of the parasitized rubens was female; the other was
male or female (the vial contained one of each and a puparium when we received it). Of
131 Gryllus grubbed in the fall of 1980, 1 male rubens and 1 male and 3 female firmus
December, 1991
Walker & Wineriter: Hosts of a Phonotactic Parasitoid 557
TABLE 1. RATES OF PARASITISMS BY ORMIA OCHRACEA IN FIELD-COLLECTED
FIELD CRICKETS, ALACHUA CO., FLA.
Fall (Sep-Dec) Other (Jan-Aug)
Species
Collecting method Year N % P Year N % P
Gryllus rubens
Sound trap 1986-7 269 64 1987 58 0
1990 106 43
Sound trap, 56 days 1986-7 61 0
1990 106 3
Pitfall 1986-7 42 5 1987 16 0
Grubbing 1980 35 3* 1981 111 0
1990 41 10
Gryllus firmus
Sound trap 1986 12 0
Pitfall 1986-7 58 0 1987 1 0
Grubbing 1980 96 4* 1981 99 0
1990 39 8
*Assuming that the puparia that were recovered and lost were 0. ochracea.
produced puparia. The puparia were lost, but subsequent results (fall 1990) suggested
that some, perhaps most, were 0. ochracea. Grubbing was resumed in fall 1990, and
0. ochracea emerged from 4 rubens (1 of 11 males and 3 of 30 females), and from 3
firmus (1 of 18 males and 2 of 21 females). Three rubens and one firmus produced
puparia of a conopid fly, Stylogaster biannulata (Say).
Exposure of lab-reared males. Of the 23 G. rubens males that were recovered and
assayed, 0 of 10 muted males and 7 of 13 nonmuted males (54%) were parasitized. (Ten
males escaped and 15 died prior to 10 days and were not dissected.) A chi square test
of the hypothesis that rates of parasitism for muted and nonmuted males were equal
yielded a P of <0.025. The level of parasitism for nonmuted males was similar to the
79% and 23% that Cade (1979, 1984) reported for calling male G. integer in central Texas.
In Alachua County, only a small proportion of G. rubens males call in the fall (TJW,
unpublished). This could account for low levels of parasitism among field-collected males.
On the other hand, the lab-reared males were apparently ready callers-as suggested
by the contrast in rates of parasitism between muted and nonmuted males and as
indicated by nonmuted males calling after exposure in the field.
Further discussion. Unlike Cade, who found only males of G. integer parasitized by
0. ochracea, we found females of G. rubens and G. firmus parasitized at rates indistin-
guishable from those of males. This was true not only for specimens collected at sound
traps, where host finding apparently was largely by common attraction to the synthesized
rubens call, but also for specimens collected at pitfalls and by grubbing. For example,
in the fall 1990 grubbing sample, 1 of 11 rubens females and 2 of 21 firms females.
Perhaps flies larviposit in response to rubens songs, but the larvae attack any Gryllus
they contact. If this be so, G. rubens should be more heavily parasitized than G. firmus.
Our data that address this issue fairly are in this direction but not significantly so. In
the 1990 grubbing samples, 4 of 41 rubens and 3 of 39firmus were parasitized. In the
pitfall samples, the corresponding figures were 2 of 40 and 0 of 58.
The levels of natural parasitism of G. rubens and G. firmus that we found are so
low that the abundance of 0. ochracea at rubens-baited sound traps in the fall is surpris-
ing. Two explanations seem worth discussing.
558 Florida Entomologist 74(4) December, 1991
(1) One or more host species other than G. rubens and G. firmus produce large
numbers of 0. ochracea. Candidate alternative hosts would be crickets that are abundant,
large enough to mature Ormia, and have a call known or likely to be attractive to O.
ochracea. Only two local species meet these requirements: S. borellii and Orocharis
luteolira Walker. The latter is arboreal and will be difficult to study (Walker 1969). For
the former species and the related S. vicinus, there are many data. We and others have
held thousands of both S. borellii and S. vicinus Scudder that were collected at sound
traps in Alachua County. The only instances of parasitism by 0. ochracea have been
three sound-trapped S. borellii. Because larvipositing 0. ochracea females are occasion-
ally attracted to S. borellii sound traps, these instances may be an artifact of the
collecting technique-as shown above for sound-trapped G. rubens. We tried manually
parasitizing S. vicinus and S. abbreviatus Scudder (n= 50 and 20) but produced no
pupae. Dissection of individuals that died showed developing larvae in S. abbreviatus
but none in S. vicinus (unpublished).
(2) Numbers of G. rubens in the fall are so great that low parasitism levels are
enough to produce an abundance of 0. ochracea. This is supported by the numbers of
G. rubens flying to sound traps. Walker (1986) reported an annual average catch of
8,209 during a 3 year study with approximately 72% being trapped during August to
November. Furthermore, only the long-winged morph of rubens is caught at sound
traps, and by several techniques Walker (1987) estimated that fewer than 50% of wild
rubens are long winged. Thus fall numbers of rubens may be great enough to produce
large numbers of 0. ochracea even at low levels of parasitism.
Either or both of these explanations could account for the abundance of 0. ochracea
in the fall. The second is surely part of the answer-and the low percentage parasitism
concurs with the observation that wild G. rubens males (unlike the reared, experimentally
exposed males) do relatively little calling in the fall.
ACKNOWLEDGMENTS
We thank Will Hudson for running the pitfall traps, Dong Ngo and P. M. Choate
for doing most of the grubbing, and J. P. Parkman for bringing us two of the parasitized
S. borellii. J. H. Frank and John Amoroso constructively criticized the manuscript. This
is Florida Agricultural Experiment Station Journal Series No. R-01483.
REFERENCES CITED
BURK, T. 1982. Evolutionary significance of predation on sexually signalling males.
Florida Entomol. 65: 90-104.
CADE, W. 1975. Acoustically orienting parasitoids: Fly phonotaxis to cricket song.
Science 190: 1312-1313.
- 1979. The evolution of alternative male reproductive strategies in field crickets.
Pp. 343-379 in M. S. Blum and N. A. Blum [eds.], Sexual selection and reproduc-
tive competition in insects. Academic Press New York.
1984. Effects of fly parasitoids on nightly duration of calling in field crickets.
Canadian J. Zool. 62: 226-228.
FOWLER, H. G., AND J. N. KOCHALKA. 1985. New records of Euphasiopteryx depleta
(Diptera: Tachinidae) from Paraguay: attraction to broadcast calls of Scapteriscus
acletus (Orthoptera: Gryllotalpidae). Florida Entomol. 68: 225-226.
LAWRENCE, K. 0. 1982. A linear pitfall trap for mole crickets and other soil arthropods.
Florida Entomol. 65: 376-377.
MANGOLD, J. R. 1978. Attraction of Euphasiopteryx ochracea, Corethrella sp. and
gryllids to broadcast songs of the southern mole cricket. Florida Entomol. 61:
57-61.
Galliart & Shaw: Paired Male Choursing
NICKLE, D. A. 1992. Scapteriscus borellii Giglio-Tos: the correct species name for the
Southern Mole Cricket in southeastern United States (Orthoptera: Gryllotal-
pidae). Proc. Entomol. Soc. Washington 94: (in press).
SABROSKY, C. W. 1953a. Taxonomy and host relations of the tribe Ormiini in the
western hemisphere. Proc. Entomol. Soc. Washington 55: 167-183.
1953b. Taxonomy and host relations of the tribe Ormiini in the western hemis-
phere, II. Proc. Entomol. Soc. Washington 55: 289-305.
WALKER, T. J. 1969. Systematics and acoustic behavior of United States crickets of
the genus Orocharis (Orthoptera: Gryllidae). Ann. Entomol. Soc. America 62:
752-762.
S1986. Monitoring the flights of field crickets (Gryllus spp.) and a tachinid fly
(Euphasiopteryx ochracea) in north Florida. Florida Entomol. 69: 678-685.
- 1987. Wing dimorphism in Gryllus rubens (Orthoptera: Gryllidae). Ann. En-
tomol. Soc. America 80: 547-560.
1989. A live trap for monitoring Euphasiopteryx spp. and tests with E. ochracea
(Diptera: Tachinidae). Florida Entomol. 72: 314-319.
WINERITER, S. A., AND T. J. WALKER. 1990. Rearing phonotactic parasitoid flies
(Diptera: Tachinidae: Ormia spp.) Entomophaga 35: 621-632.
EFFECT OF INTERMALE DISTANCE AND FEMALE
PRESENCE ON THE NATURE OF CHORUSING BY
PAIRED AMBLYCORYPHA PARVIPENNIS
(ORTHOPTERA: TETTIGONIIDAE) MALES
PATRICK L. GALLIART AND KENNETH C. SHAW
Department of Zoology & Genetics
Iowa State University
Ames, IA 50011
ABSTRACT
Pairs of chorusing Amblycorypha parvipennis Stal males alternate the production
of 3-5 s long phrases with frequent partial overlap of phrases. To determine the effect
of intermale distance and the presence of a sexually receptive, sound-producing ("tick-
ing") female on the nature of paired male chorusing, pairs of males were recorded, with
and without the presence of a ticking female placed midway between the caged males,
chorusing 3.3 m and 40 cm apart. The presence of a ticking female elicited a shortening
of intervals between phrases (increase in phrase rate) and an increase in the extent of
phrase overlap. In contrast, shortening the distance between males in the absence of a
ticking female resulted in a lengthening of phrase interval (decrease in phrase rate) and
a reduction in phrase overlap. Possible proximate and ultimate causes of these chorusing
changes are discussed.
RESUME
Pares de machos de Amblycorypha parvipennis Stal, cantando en coro, alternan la
production de frases de 3-5 segundos de duracion, con la frequent sobreposicion parcial
de las frases. Para determinar el efecto de la distancia entire machos y la presencia de
una hembra sexualmente receptive y productora de sonido en la naturaleza del par de
machos cantando en coro, los pares de machos fueron grabados, con y sin la presencia
560 Florida Entomologist 74(4) December, 1991
de la hembra la cual estaba situada a 3.3m y 40 cm de distancia de los machos en jaula.
La presencia de la hembra produjo un acortamiento entire los intervalos entire frases
(incremento en la production de frases) y un incremento en la sobreposicion de estos.
En contrast, al acortar la distancia entire machos y en la ausencia de una hembra,
result en la extension del intervalo de frases (la production de frases decrecio) y en
una reduction en la sobreposicion de las frases. Se discuten las causes inmediatas y
ultimas para estos cambios en los coros.
Available evidence suggests that both inter- and intrasexual selection are important
in the evolution of chorusing between and among male singing Orthoptera (see reviews
by Alexander 1975, Otte 1977, Greenfield & Shaw 1983, Greenfield 1990). Male compe-
tition for females is expressed in uniform spacing of singing males resulting from move-
ments away from and/or towards the songs of adjacent males (Campbell & Shipp 1979,
Thiele & Bailey 1980, Bailey & Thiele 1983, Latimer & Schatral 1986, Latimer & Sippell
1987). Associated with spacing dynamics, singing orthopteran males may alter their
songs when another singing conspecific male moves closer (Alexander 1957, 1961, Fever
1977, Otte 1977, Meixner & Shaw 1986, Shaw 1968). If uniform spacing serves to reduce
male competition for females, then a reduction in distance between neighboring males
should represent an increased threat to both males' abilities to acquire mates. An obvious
reaction to such a threat would be to increase energy output by increasing sound output
and/or entering into physical combat. Males of many field cricket species may increase
length of chirps, reduce intervals between chirps and/or increase sound level of chirps
(Alexander 1957, 1961). This also is true of males of at least some species of the
pseudophylline katydid, Pterophylla (Shaw 1968, Shaw & Galliart 1987, Barrientos
1988). Males of several species of the concocephaline genus, Orchelimum, increase the
rate, length, and/or intensity of the "tick" component of their calling sounds when a
conspecific intruder approaches (Feaver 1977).
In contrast, a study utilizing males of the Australian conocephaline katydid Mygalop-
sis marki Bailey suggests that males of this species may reduce the number of pulses/
phrase and reduce phrase rate by increasing intervals between phrases in response to
a reduction in distance between neighboring males. Reduction in distance between an
apparent intruder and a territorial male was simulated by increasing the sound level of
playbacks of a M. marki male song and recording the response of territorial males in
the field (Dadour 1989).
Since females of most species of singing Orthoptera move toward males that are
1 m or more from other singing males, male acoustic interaction in competition over a
nearby female is seldom seen. Such encounters have been reported between cricket
males in terraria (e.g., Alexander 1961, Burke 1983), but there has been no report that
the nature of sounds or the acoustic interaction differs from that when a female is
present. In species of the subfamily Phaneropterinae, females produce simple sounds
in response to male calls and the males move to the female (Spooner 1968). This method
of bringing the sexes together should increase the opportunity for male-male acoustic
interaction at close range. Although Spooner (1968) indicates that phaneropterine males
sing differently in groups than in isolation, we know of no investigation of the effect of
change in intermale distance or the presence of a female on the nature of acoustic
interaction of males in any species of this subfamily.
The phaneropterine katydid, Amblycorypha parvipennis StAl is unique among
chorusing Orthoptera in that males alternate overlapped phrases and, where phrases
overlap, phrase subunits (phonatomes) are synchronized (Shaw et al. 1990) (Fig. 1).
This study asks whether the nature of sound production and acoustic interaction of
paired males are affected by moving the males closer together in the absence and
presence of a sound-producing female.
__ __ __~\
___~
Galliart & Shaw: Paired Male Choursing 561
A -a- PHRASE
I I-...l'U TII4 T -I Ii.: Hi. I$ t I IpINIINll Illill ll l )I III 1 ll. | IIIll tIl T9 tlh IIm II(til l
b PHONA TOML
B
2 1 q 111 to m 11 jil 111 M 111 1 1 11A11 1 14 1W if W W if 1 *
__ _ _- --.b -----
0 8 16 24
TIME IN ';EC
Fig. 1. Oscillographs of chorusing by paired A. parvipennis males. A the initial
part of the phrase of each male overlaps the latter part of the phrase of the other male.
B in this selection, katydid 2 overlaps katydid 1 but 1 does not overlap 2. a indicates
the time that the phrase of 1 overlaps the phrase of 2. b indicates the time that the
phrase of 2 overlaps the phrase of 1.
MATERIALS AND METHODS
Subjects and Housing
A. parvipennis males and females were collected at night using flashlights during
June and July, 1986-1989. Most virgin females were collected during the first week of
singing. Males were collected as needed throughout the testing period. Specimens were
marked with fingernail polish. Males were isolated in 10x10x17 cm wire screen cages
and females were housed together in 34x34x31 cm wire screen and wood cages. The
insects were housed in a laboratory maintained on a 14L:10D light schedule at 24-250C.
The insects were fed leaves of horsemint (Mentha longifolia (L)). or wild grape (Vitis
sp.), and water was provided in cotton-capped vials.
The Effect of Intermale Distance and Female
Presence on Paired Male Chorusing
In order to investigate the effect of intermale distance and female presence on male
chorusing, we compared 10 min recordings of paired males singing under four different
conditions: 1) two caged males 3.3 m apart (within the range of common, field nearest
neighbor distances [Shaw et al. 1981]), 2) males 40 cm apart (a proximity which should
elicit male-male competition for space), 3) males 3.3 m apart with a sexually receptive
(ticking) female placed midway between the two males, and 4) males 40 cm apart with
a ticking female midway between the males. The recordings were made using two
unidirectional dynamic microphones (GC Electronics, #30-2374), each placed 6 cm from
the cage of a singing male, and a Sony TC-6300, 2-channel tape recorder. The temporal
parameters of each male's song and the song phrase phase relationships of pairs of
chorusing males were determined using a Commodore 128 computer in conjunction with
a computer interface and software designed for this analysis.
The data used in this paper are taken from studies originally designed to determine
which acoustic and other factors affect mating success and male competition. Data for
conditions 1) and 3) were taken from 114 males used in 57 two-choice discrimination
Florida Entomologist 74(4)
trials run in 1986 and 1987. In these trials males were recorded chorusing 3.3 m apart
in the absence of a female and with a ticking female placed midway between the two
caged males. After the recordings, the males and female were released in order to
determine which male eventually mated with the female (Galliart & Shaw 1991). In
1988, we obtained data for conditions 1) and 2) during male competition trials. In 22
trials, 44 males were recorded chorusing at 3.3 m and 40 cm prior to release from the
cages to determine which male stayed and which left the immediate area (unpublished
data). In 1989, we returned to two-choice discrimination trials during which we obtained
data from 66 males for conditions 1), 3) and 4). These 33 trials were the same as in 1986
and 1987 except that pairs of males were recorded at 40 cm apart following a recording
at 3.3 m and then released (unpublished data).
Data Analysis
Because of the difficulty in collecting specimens, especially females, males and females
were used in more than one trial. Males were used in up to three different trials (85%
males used once, 10% used twice, 5% used three times); however, reused males were
never matched with the same male twice. As discussed later, this source of males does
not affect the experimental error values used in performing tests of probability. Females
were used up to five times but, unlike two-choice discrimination tests, using females
more than once as aa source of ticking sounds should have little effect on the results
analyzed in this paper.
The 10-min. segments of paired males chorusing under each condition were examined
in relation to temporal sound parameters (phrase number, phrase length, phrase interval,
phrase period [phrase length + phrase interval], total sound produced in 10 min [phrase
number x phrase length]) and nature of phrase overlap (number of phrase overlaps,
mean overlap time and total overlap in 10 min [number of phrases overlapped x mean
time of overlap]). Phrase overlap for each katydid was measured from the time that a
katydid initiated its phrase during the phrase of another katydid until the other katydid
terminated its phrase (Fig. 1). Phrase overlap does not include the latter portion of a
katydid's phrase that is overlapped by the initiation of the other katydid's phrase.
In order to combine data from different years and compare data across years, we
performed an analysis of combined experiments (Cochran & Cox 1957) (see Table 1 for
example). This analysis treats the mean value for each condition during each year as a
different measurement (e.g. condition 1 is treated as four measurements, one each year,
not as 224 measurements over four years). Therefore the experimental error values
used in all statistical tests involve between year variation rather than variation within
any condition during one year. The four conditions were considered the treatments. The
treatments were regarded as a factorial combination of the factors of intermale distance
TABLE 1. Use of analysis of combined experiments to analyze the effect of intermale
distance and female presence on the phrase intervals of paired chorusing
males.
Source df Mean Square P > F
Year 3 0.12306 0.0022
Intermale Distance 1 0.04079 0.0066
Female Presence 1 0.64659 0.0004
Distance*Female 1 0.12681 0.0021
Error 2 0.00027
December, 1991
Galliart & Shaw: Paired Male Choursing
and presence or absence of a female. The response variables are the acoustic parameters
listed above. This analysis allowed us to look at the main effect of female presence, the
main effect of distance and the interaction between these two factors. When the interac-
tion between female presence and distance was significant, least square means for the
appropriate singing categories were compared using two-tailed Student's t-tests. Least
square means were used to adjust means for differences among years and to enable
comparison of chorusing at 40 cm in the absence and presence of a female, two conditions
which were recorded during different years.
RESULTS
The presence of a female had an excitatory effect on male singing behavior; following
introduction of a ticking female, males increased their phrase rate. At both intermale
chorusing distances, the phrase period (reciprocal of phrase rate) was less when a ticking
female was present (Fig. 2; F = 68.73; df = 1,2; p = 0.014). The reduction in phrase
period was a result of a shortening of phrase interval (Fig. 3; F = 118.38; df = 1,2; p
= 0.008).
p-0.024
-- p-0.347
3.3
40 cm
INTERMALE DISTANCE
Fig. 2. The effect of intermale distance and female presence on the phrase period of
pairs of chorusing A. parvipennis males. Males were recorded in the absence (solid line)
and presence (dotted line) of a ticking female. P values from Student's t-tests.
9.6-
9.4-
9.2-
9.0-
8.8-
8.6-
8.4-
563
Florida Entomologist 74(4)
5.0-
4.8-
4.6-
4.4-
4.2-
4.0-
3.8-
3.6-
December, 1991
p-0.0 19
S\
p-0.081
3.3
40 cm
INTERMALE DISTANCE
Fig. 3. The effect of intermale distance and female presence on the phrase intervals
of pairs of chorusing A. parvpennis males. Remainder as in Fig. 2.
The effect on the nature of chorusing of moving males closer together was contingent
upon female presence. There was a significant interaction between intermale distance
and female presence for phrase interval (Fig. 3; F = 56.92; df = 1,2; p = 0.017) and
the interaction for phrase period approached significance (Fig. 2; F = 15.17; df = 1,2;
p = 0.060). In the absence of a female, the effect of moving chorusing males closer
together was opposite to that of female presence, i.e., there was an inhibitory effect
expressed as a reduction in phrase rate. Both mean phrase period (Fig. 2) and interval
(Fig. 3) increased significantly when intermale distance was reduced from 3.3 m to 40
cm. The presence of a ticking female not only elicited an increase in phrase rate of the
chorusing males, it eliminated the effect of distance reduction between chorusing males
(Figs. 2, 3).
564
Galliart & Shaw: Paired Male Choursing 565
Female presence and the reduction in intermale distance also affected the degree of
phrase overlap by chorusing males. The presence of a female resulted in an increase in
mean phrase overlap (Fig. 4; F = 2371.61; df = 1,2; p = 0.0004) and total overlap (Fig.
5; F = 45.65; df = 1,2; p = 0.021) at both intermale distances. As for phrase rate,
decreasing the distance between chorusing males in the absence of a female had effects
opposite to that of introducing a female. There was a significant interaction between
female presence and intermale distance for both phrase overlap (Fig. 4; F = 465.13; df
= 1,2; p = 0.002) and total overlap (Fig. 5; F = 18.74; df = 1,2; p = 0.049). When
males were moved closer together in the absence of a female, there was a decrease in
both mean (Fig. 4) and total (Fig. 5) phrase overlap; however, the latter difference was
not statistically significant. As in the case of phrase rate, the presence of a female
eliminated (Fig. 5) and even reversed (Fig. 4) the effect of reducing distance in the
absence of a female.
2 0
.-*p-0.020
-J
1.0-
a,
n"
1.5-
O
I 1.0
< p-0.002
0.5 -
3.3 m
40 cm
INTERMALE DISTANCE
Fig. 4. The effect of intermale distance and female presence on the extent to which
each of a pair of A. parvipennis males overlapped the phrases of their chorusing partner.
Remainder as in Fig. 2.
Florida Entomologist 74(4)
*p-0.130
90-
80-
70-
60-
50-
40-
30-
20
40 cm
INTERMALE DISTANCE
Fig. 5. The effect of intermale distance and female presence on the total period of
time (number of phrases produced in 10 min x mean phrase overlap) that a pair of A.
parvipennis males overlapped one another's phrases. Remainder as in Fig. 2.
DISCUSSION
In analyzing the nature of chorusing by pairs of katydid males, several investigators
have provided evidence that alternation is the result of one male being inhibited during
the phrase of another male (Jones 1966, Shaw 1968, Samways 1976, Latimer 1981).
Because the phrase intervals of A. parvipennis males are longer during acoustic interac-
tion than when males sing alone, Shaw et al. (1990) suggested that alternation of over-
lapped phrases results from one male being inhibited for some period during the song
phrase of another katydid. Smith (1986) showed that electronically produced imitations
falling between an A. parvipennis male's phrases increased phrase interval when com-
pared to males singing alone or when imitations occurred during a male's phrases.
Moving two A. parvipennis males closer together than 3.3 m appeared to enhance
inhibition, i.e., increased phrase interval was associated with reduced phrase overlap,
p-0.070
3.3 m
December, 1991
Galliart & Shaw: Paired Male Choursing
567
the latter resulting in more of the phrase of one katydid falling in the phrase interval
of the other katydid. This increased inhibitory effect could have been the result of the
increased intensity with which one katydid heard the sound phrases of the other.
Spooner's (1968) analysis of male-female and male-male acoustic interaction of a
number of phanopterine species suggests that inhibition may not be uncommon in the
acoustic interaction of conspecific males. In congregations of Inscudderia strigata (Scud-
der) and Amblycorypha floridana Rehn and Hebard males, phrases (lisps or buzzes)
produced by one male or a playback, resulted in a cessation of phrases by other males.
If acoustic interaction is involved in spacing dynamics, and if reduction in distance
between two conspecific males increases the competition between the males, why would
males reduce rate of sound output under the latter conditions? If, as Dadour (1989)
suggests for M. marki males, increasing phrase interval between phrases enhances a
male's ability to monitor an intruder (and, we assume, an intruder's ability to monitor
a resident), then an increase in phrase interval, as well as a reduction of phrase overlap
by A. parvipennis males, may facilitate the decision as to which male leaves the area
of confrontation.
This reduction in rate of sound output is contrasted to an increase in rate of sound
output evidenced when chorusing males were exposed to a ticking female (Fig. 2). The
presence of a female also negated the effect of reduction in distance. Thus, the importance
of competing for a sexually receptive female completely superseded the effect of male-
male competition in the absence of a female.
In addition to increasing phrase rate in the presence of a ticking female, males
increased phrase overlap. This phenomenon, which has been discussed in detail previ-
ously (Galliart & Shaw 1991), will only be summarized here. In the data utilized in this
study to compare the nature of chorusing of male pairs in the absence and presence of
a female, the male that usually succeeded in copulating with the female (the winner),
overlapped the phrases of the other male (the loser) less than vice-versa (e.g., Fig. 1B).
In fact, the total number of phrases overlapped for both katydids decreased in the
presence of a female, but the decrease was principally due to a decrease on the part of
the winner (Galliart & Shaw 1991). Thus, although the number of phrases overlapped
decreased, the mean length of phrase overlap increased suggesting that this increase
is a byproduct of males adjusting their phrase rates in an attempt to reduce the number
of phrases they overlap. A preliminary analysis of three paired interactions has indicated
that a male's extent of phrase overlap is greater when his chorusing partner does not
overlap the male's previous phrase than when the chorusing partner does overlap it
(unpublished data) (e.g., compare extent of phrase overlap by katydid 2 in Figs. 1A and
1B).
The question remains as to why males of some species increase sound output and
others decrease it when distance between singing males is decreased. A number of
factors could be involved. The importance of specific singing sites may determine the
degree of energy expended. Field crickets occupy a burrow or equivalent which they
defend and which may be important to success in mating with a female. In Orchelimum
spp., the position within a resource patch may be important in controlling female access
to the patch (Feaver 1977, 1983). In contrast, in other species, specific areas may be
important because of an increased chance of encountering females, but occupation of
specific sites within these areas may not be important. For example, in A. parvipennis,
male and female density is related to density of preferred host plant (Shaw et al. 1981).
However, because females tick in response to male sounds and two or more males may
move to a ticking female, specific sites within a resource patch may not be important
to defend.
Also, a mechanism of intermale communication which involves a reduction in energy
output in the absence of a female may be favored by selection in order that energy is
568 Florida Entomologist 74(4) December, 1991
conserved for male-male and male-female competitive communication in the presence
of a sexually receptive female. In the presence of a ticking female, A. parvipennis males
encounter one another physically and acoustically and male-male and male-female encoun-
ters may last for many hours. During one night's observation, we intermittently observed
three females, each being courted by 4-5 males, from our initial observation (between
10 P.M. and 12 midnight) until dawn. None of the females mated (Shaw et al., unpublished
observations) before dawn. In the laboratory, we have run 114 mate choice trials with
two males and one virgin female and, in 24 of these trials, copulation had not occurred
in the 4-hour limit given to the trials. During the mate choice trials, males lost up to
12% of their body weight (Galliart & Shaw, unpublished observations).
With the emphasis on male-male and male-female strategies and the application of
game and economic theory to an understanding of animal reproductive behavior, (Krebs
& Davies 1984; Maynard Smith 1989) changes in acoustic interaction of conspecific males
with change in intermale distance and presence of absence of females should be of special
interest to animal behaviorists. The ease with which sounds can be recorded and analyzed
make such analyses relatively easy. In some species, quantitative and/or qualitative
changes, termed aggressive and courtship (Alexander 1967, Otte 1977) are obvious. This
and Dadour's (1989) study have shown that interesting and significant changes in calling
songs, which may not be obvious to the unaided ear, may be taking place in many species
not characterized by the production of obvious aggressive and courtship songs.
ACKNOWLEDGMENT
We wish to thank Dr. Paul Hinz, Department of Statistics, Iowa State University,
for recommending and running the analysis of combined experiments. This research
was partially funded by grants from the Iowa Science Foundation.
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Florida Entomologist 74(4)
IMPROVEMENT IN EFFICACY OF GIBBERELLIC ACID
TREATMENTS IN REDUCING SUSCEPTIBILITY OF
GRAPEFRUIT TO ATTACK BY CARIBBEAN FRUIT FLY
P. D. GREANY
Insect Attractants, Behavior,
and Basic Biology Research Laboratory
Agricultural Research Service, USDA
Gainesville, FL 32608
R. E. MCDONALD AND W. J. SCHROEDER
U.S. Horticultural Research Laboratory
Agricultural Research Service, USDA, Orlando, FL 32803
P. E. SHAW
Citrus and Subtropical Products Research Laboratory
Agricultural Research Service, USDA, Winter Haven, FL 33880
ABSTRACT
Efficacy of the plant growth regulator, gibberellic acid (GA3), in reducing 'Marsh'
grapefruit susceptibility to Anastrepha suspense was increased through use of the
surfactant, L-77 Silwet. In addition, it was found that the duration of protection of
grapefruit could be extended by use of two consecutive GA3 treatments a month apart.
Corresponding effects on peel color and resistance to mechanical puncture were noted,
but peel oil content was not affected by treatment and declined at the same rate as that
of untreated fruit. The implications of these results in the further development of the
fly-free concept and for fruit fly control are discussed.
RESUME
Se demuestra que el uso de un surfactante mejorado (L-77 Silwet) incrementa la
eficacia del regulador de crecimiento Acido Giberelico (AG) en reducir la susceptibilidad
de toronjas de la variedad "Marsh" al ataque de Anastrepha suspense. Ademas se
demuestra que dos aplicaciones consecutivas (un mes de intervalo entire aplicaciones)
permiten extender el period durante el cual las toronjas resisten el ataque de moscas.
Los tratamientos con AG tambien ocasionaron cambios en el color de la cascara del fruto
e incrementaron la resistencia mecanica a la penetracion de una aguja. Sin embrago, el
contenido de aceites no se vio afectado. Estos descubrimientos, una vez confirmados en
pruebas de campo, pueden tenener implicaciones importantes en el desarrollo de metodos
alternatives de control de moscas de la fruta. En especial se consider que se lograra
apoyar el desarrollo del concept de "areas libres de moscas".
As citrus fruit become senescent, they become more susceptible to attack by tephritid
fruit flies (Greany et al. 1983, 1985, Greany 1989). However, the natural plant growth
regulator, gibberellic acid (GA3), has proven useful in delaying senescence-related
changes in citrus fruit peel without interfering with internal ripening (Coggins 1973,
Ali Dinar et al. 1976, Ferguson et al. 1982, Lima & Davies 1984, McDonald et al. 1987).
Prior studies by Greany et al. (1987), McDonald et al. (1987), and Rossler & Greany
(1990) have shown that by treating grapefruit and oranges with GA3 prior to fruit
colorbreak, it is possible to significantly reduce their susceptibility to attack by the
570
December, 1991
Greany et al.: Reducing Fruit Fly Susceptibility
Caribbean fruit fly, Anastrepha suspense (Loew), and the Mediterranean fruit fly,
Ceratitis capitata (Wiedemann).
If these findings can be confirmed in field tests, they have considerable potential
importance for the control of fruit flies in citrus. Thus, it may be possible to use GA3
treatments in addition to, or in lieu of, other pre- and postharvest treatments intended
to assure the absence of fruit flies from citrus fruit (McDonald et al. 1988). Because of
the abolition of postharvest fumigation with ethylene dibromide and the unpopularity
of available postharvest cold treatments with fruit growers, packers, and shippers, the
use of the fly-free concept for quarantine purposes is gaining in popularity in Florida
and elsewhere (Riherd 1991). Currently, about 50% of Florida grapefruit are certified
for quarantine purposes under one of the fly-free protocol provisions (i.e., without
postharvest treatment (C. Riherd, Division of Plant Industry, Florida Dept. of Agricul-
ture, personal communication). Consequently, GA3 treatments could provide a valuable
tool to reduce the likelihood of fruit fly infestations in citrus fruit. GA3 treatments
already are being used in some areas to extend the citrus harvest season and to reduce
postharvest fruit susceptibility to decay and mechanical injury without affecting internal
ripening (Greenberg et al. 1987, and references therein). This use is approved by the
U.S. Environmental Protection Agency. To extend this approach to assist in management
of fruit flies as well as postharvest decay would provide a dual benefit from a single
therapy, making GA3 use even more cost-effective. Should prophylactic malathion-pro-
tein hydrolyzate bait sprays (an important tool in current fruit fly management and
abatement strateiges) be banned, development of alternative methods for protection of
fruit prior to harvest will become increasingly important.
Toward improving GA3 efficacy, and to reduce the cost of treatment, Greenberg et
al. (1984, 1987) found that a nonionic organo-silicone copolymer surfactant, L-77 Silwet
(Union Carbide) improved the uptake of GA3 by citrus fruit peel as compared with use
of the wetting agent Triton B 1956 (Rohm & Haas). This allowed about a five-fold
reduction in the concentration of GA3 needed to achieve a given level of effectiveness,
with commensurate cost reduction.
The present studies were conducted to determine whether use of L-77 under Florida
conditions also would allow a reduction in the amount of GA3 required to delay senescence
of grapefruit and thereby sustain fruit resistance to A. suspense. In addition, we sought
to determine whether repeated GA3 treatments a month apart would allow an even
longer refractory period.
MATERIALS AND METHODS
Studies were conducted for two consecutive seasons in a citrus grove located adjacent
to the Merritt Island National Wildlife Refuge, Merritt Island, Florida. 'Marsh' grape-
fruit trees (Citrus paradise Macf.) on rough lemon rootstock (C. jambhiri Lush.) were
employed for the tests. The trees were about 16 years old and about 7 meters in height.
Fruit Treatment 1987/88 Season
A comparison was made of use of the wetting agent previously used with GAs
(McDonald et al. 1987), Triton X-100, vs. L-77.
Triton X-100 group: Six trees each received one of three treatments replicated two
times. The fruit were green in color when treated on 15 September 1987 with 0, 20, or
50 pl/liter (0, 20 or 50 ppm) GA3 (Pro-Gibb, Abbott Labs) along with 0.1% (v/v) of a
wetting agent (Triton X-100). The trees were sprayed until run-off, or with about 25
liters per tree.
Florida Entomologist 74(4)
L-77 group: Six similar trees in the same grove also were sprayed with 0, 5, or 10 Il/liter
(0, 5, or 10 ppm) GA3 along with 0.1% (v/v) of the surfactant L-77 Silwet (dimethyl
polysiloxane modified with alkylene oxide). Two additional trees were left unsprayed
as controls.
Fruit Treatment 1988/89 Season
Only one concentration of GA3 was employed for all GA3 treatments, 10 pl/liter (10
ppm) GA3, and only L-77 was used, based upon encouraging results from the previous
season. The concentration of L-77 was varied to determine the minimum effective con-
centration because both L-77 and GAs are expensive. Eight trees were sprayed to run-off
on 13 September 1988 with aqueous solutions of 0.025% or 0.1% L-77, along with 10
ppm GA3 (4 replicates per treatment). Of these, on October 17, 2 trees were re-sprayed
with 0.025% L-77 and 10 ppm GA3 and 2 with 0.1% L-77 and 10 ppm GA3. Two trees
were left unsprayed for use as controls.
Peel Puncture Resistance and Color Measurement
At monthly intervals after treatment, fruit were harvested for analyses. For peel
firmness, eight fruit, evenly spaced around each tree, were picked. The firmness of the
rind tissue was evaluated in terms of resistance of the rind to puncture, using an Instron
Food Testing System (Model 1132) pressure testing device, set to apply a constant
weight of 10 kg at a constant rate of travel of 5 cm/min to a 1.0 mm diam. cylindrical
tip. Four punctures were made on each test fruit at evenly spaced locations in the
equatorial region of the fruit, and the force required to puncture the peel recorded.
Color was measured, after washing the fruit with water only, on the same 8 randomly
selected fruit with a Hunterlab Colorimeter (Model D25-9). Three color readings were
made on each test fruit at evenly spaced locations in the equatorial region of the fruit.
A Hunter "a/b" ratio of-0.10 was considered as the minimum yellow color for grapefruit
to be used commercially. An increasing Hunter "a/b" value is indicative of a color change
from green to red, concomitant with a change from blue to yellow.
Peel Oil Measurement
The effect of GA3 treatment on peel oil content was measured each season, using
the techniques described by Wilson et al. (1990).
Fruit Fly Susceptibility Bioassays 1987/88 Season
The methods described by Greany et al. (1987) were employed for laboratory exposure
of treated and control fruit in choice tests. Each month beginning in November 1987,
fruit were harvested from the circumference of the trees (to avoid a position effect),
washed gently after harvest with cool tap water, and allowed to air dry. Two fruit of
each treatment type were arranged on a platform suspended about 15 cm below the top
of 36 x 36 x 46 cm exposure cages. This test was replicated six times each month. We
used a ratio of 1 fruit to 4 gravid 10-to-15 day old A. suspense females reared on artificial
media in the laboratory (not reared on fruit). The exposure period was 3 days. Food
(enzymatic yeast hydrolyzate and sucrose) and water were provided. Ambient conditions
were as follows: fluorescent light (500 lux) with a 12:12 L:D photoperiodic regime, about
25C and 50% RH. After exposure, fruit were placed in individual ventilated containers
on a pupation medium (vermiculite) for 2 months to determine the success in fly devel-
opment).
Only fruit treated with GA3 and L-77 (not those treated with GA3 plus Triton X-100),
plus controls, were bioassayed. This was because we had already bioassayed fruit treated
December, 1991
Greany et al.: Reducing Fruit Fly Susceptibility
with GA3 plus Triton X-100 at 20 and 50 ppm (Greany et al. 1987). Direct comparisons
between L-77 and Triton X-100 were made only on the basis of observed effects on peel
color and puncture resistance, rather than fruit fly susceptibility.
Fruit Fly Susceptibility Bioassays 1988/89 Season
Fruit treated with GA3 vs. controls were exposed and then held for progeny devel-
opment as indicated above, except that only 1 gravid A. suspense female was provided
per fruit during the exposure period, to allow a less extreme forced-infestation situation.
Again, two fruit of each treatment type were exposed together each month, beginning
in October 1988. Ten cages (replicates) were set up each month.
Statistical Analyses
Mean success in fly development (average number of flies, including unemerged
puparia, produced per fruit) among treatments was evaluated by using ANOVA and
Duncan's new multiple range test.
RESULTS
1987/88 Season
All GA3 treatments resulted in retained peel puncture resistance, as compared to
the control fruit (Fig. 1). There was no apparent effect of either Triton X-100 or L-77
employed with 0 ppm GA3. The effect of treatment was greatest using 10 ppm GA3 with
L-77, even greater than from 50 ppm GA3 with Triton X-100. Fruit treated with the 10
ppm GA3/L-77 formulation were about 55% firmer than control fruit from January
through April. The effect from only 5 ppm GA3 with L-77 was nearly as pronounced as
from 20 ppm GA3 with Triton X-100, at least until January, when the 5 ppm/L-77 fruit
began to soften more rapidly than the 20 ppm/Triton X-100 fruit.
The GA3-treated fruit also remained greater in color compared to the control fruit,
as indicated by objective color readings (Fig. 2). When the fruit were left on the tree,
control fruit reached a yellowish-green considered to be minimally acceptable from a
commercial standpoint by the first of December (-0.1 Hunter "a/b"). Fruit treated with
the 5 ppm GA3/L-77 combination reached that point in late December. Fruit treated
with 50 ppm GAs/Triton X-100 reached this color standard by early January. Those
treated with 10 ppm GA3/L-77 or 20 ppm GA3/Triton X-100 degreened last, in mid-
January. Thus, use of L-77 again allowed a pronounced effect of GA3 even at a reduced
concentration, compared with the dose required using Triton X-100.
Data on the effect of GA3 treatments on peel oil abundance during the 1987/88 season
were published separately ( Wilson et al. 1990). Generally, they noted a small but
significant reduction in the rate of loss of peel oil in whole fruit as a result of treatment
with 20 or 50 ppm GA3 plus Triton X-100 as compared with control fruit. Measurements
of peel oil effects of GA3 plus L-77 were not made during this season.
Fruit fly susceptibility was significantly affected in a dose-dependent fashion by
treating the fruit with GAs (Fig. 3). As indicated earlier, only fruit involving GA3 plus
L-77 were used (not GAs plus Triton X-100) for these tests. Through December 1987,
no significant differences were noted among the treatments, but thereafter a trend of
reduced susceptibility was noted, especially in March 1988. It appeared that the 5 ppm
GA3 dose was not reliable in reducing fruit susceptibility after December. The degree
of benefit achieved from 10 ppm GA3 with L-77 was in keeping with that formerly
observed using 50 ppm GAs with Triton X-100 (Greany et al. 1987).
Florida Entomologist 74(4)
8.0
7.0
6.0
5.0
4.0
3.0
S
8.0-
7.0
6.0
5.0
4.0
3.0
I I I I
SEPT OCT NOV DEC
1987-1
I I I I
JAN FEB MAR APR
I I I
JAN FEB MAR APR
Fig. 1. Effect of GA3 treatment and surfactant on grapefruit peel puncture resist-
ance-1987/88 season. Note: puncture resistance values reported here are ca. 10x higher
than those reported in previous years by McDonald et al. 1987, and McDonald et al.
1988. This is due to an error in calibration of test equipment in prior years. Vertical
bars represent std. errors.
-*-- ntreaed .Y
---- Untreated
---0--- Oppm GAL-77
o 5 ppm GA L-77
- -A- 10 ppm GA L77
I I IV
EPT OCT NOV DEC
- Untreated
--- 0- ppm GA Triton
S 20 ppm GA Triton
- -A- 50 ppm GA Triton
December, 1991
-, H-;i--
Greany et al.: Reducing Fruit Fly Susceptibility
0.1
0
-0.1
0 -0.2
-0.3
-0.4
-0.5
-0.6
-0.7
SEPT OCT NOV DEC JAN FEB MAR APR
MONTH
0.1 -
0
-0.1-
-0.2
-0.3-
-0.4
-0.5
-0.4
-r 7 -
SEPT OCT NOV DEC JAN FEB MAR APR
SEPT OCT NOV DEC JAN FEB MAR APR
1987-88
Fig. 2. Effect of GA3 treatment and surfactant on grapefruit peel color-1987/88
season. The dotted line at Hunter "a/b" ratio-.1 represents the minimal amount of
degreening required for marketability. A low a/b ratio corresponds to a green peel color.
Vertical bars represent std. errors.
---- UNTREATED
--- -- 0 ppm GA Triton
...... .. 20 ppm GA Triton
- -- 50 ppm GA Triton
.m
576
25
Florida Entomologist 74(4)
E UNTR. CONTROL
U 0 PPM GA + L77
B 5 PPM GR + L77
A
ll 10 PPM GR + L77 I
NOV '87 DEC '87 JRN '88 FEB '88 MAR '88
MONTH
Fig. 3. Reduction in susceptibility of grapefruit to attack by A. suspense through
GA3 treatment. Vertical bars represent std. errors. Bars annotated with different letters
are significantly different at the 5% level by Duncan's new multiple range test.
1988/89 Season
Because of the success in use of reduced concentrations of GA3 when used with the
surfactant L-77 during the 1987/88 season, only L-77 was used during the 1988/89 season.
Again, a dose-dependent reduction in peel softening due to use of GA3 was noted (Fig.
4). The degree to which softening was retarded was a function of the concentration of
L-77 (0.025% vs. 0.1%) and the number of applications of the GA3/L-77 formulation. In
general, 0.1% L-77 proved superior to 0.025%, even when two applications were made
a month apart using 0.025% L-77. Fruit receiving two applications of 10 ppm GA3 with
0.1% L-77 remained quite puncture-resistant, even until April.
This trend also was exhibited for effects on fruit color (Fig. 5). Untreated fruit
reached the -0.1 Hunter "a/b" ratio by mid-December, and treated fruit required 4-6
additional weeks to reach this point. Fruit treated with consecutive applications of 10
ppm GA3 with 0.1% L-77 reached -0.1 Hunter "a/b" only at the beginning of April, and
would have required ethylene degreening to achieve marketability prior to that time.
Fruit receiving only one application of 10 ppm GA3 with 0.1% L-77 reached marketable
color status March 1st.
Peel oil abundance was not affected significantly by GA3 treatment during the 1988/89
season (data now shown). This is in contrast with the small, but significant, effect of
GA3 applied with Triton X-100 during the preceding season (Wilson et al. 1990), and
may be attributable to the use of L-77 instead of Triton X-100.
Fruit fly susceptibility was dramatically affected by use of just 10 ppm GA3 with
0.1% L-77, especially when fruit were treated in both September and again in October
(Fig. 6). This was most apparent in April 1989, when the fruit treated twice using 10
December, 1991
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