Title: Florida Entomologist
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098813/00069
 Material Information
Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1990
Copyright Date: 1917
Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
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Bibliographic ID: UF00098813
Volume ID: VID00069
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: Open Access

Full Text

(ISSN 0015-4040)


(An International Journal for the Americas)

Volume 73, No. 2 June, 1990


Announcement 73rd Annual Meeting ..................................... .............. i
Announcement Forum Section ............................................. ............... i

Research Reports
STANSLY, P. A., AND W. SANCHEZ R.-Biology and Oviposition Behavior of
Cydia fabivora (Lepidoptera: Tortricidae) in Soybean on Ecuador's Coastal
P lain ............................................... ........................................ 219
HOWARD, F. W.-Population Suppression of Mahogany Webworm, Macalla
thyrsisalis (Lepidoptera: Pyralidae) With Natural Products ................... 225
LYE, B.-H., C. W. McCoY, AND J. FOJTIK-Effect of Copper on the Residual
Efficacy ofAcaricides and Population Dynamics of Citrus Rust Mite (Acari:
Eriophyidae) .............................................................................. 230
COATS, S. A., AND M. A. ISMAIL-Ovicidal Effects of Gamma Radiation on Eggs
of the Fuller Rose Beetle, Pantomorus cervinus (Coleoptera: Curculionidae)
VON WINDEGUTH, D. L., AND W. P. GOULD-Gamma Irradiation Followed by
Cold Storage as a Quarantine Treatment for Florida Grapefruit Infested
W ith Caribbean Fruit Fly ............................................................. 242
SNIDER, R. J.-A New Species of Ptenothrix and Records From the Southeastern
United States (Dicyrtomidae: Collembola) .......................................... 248
CASANI, J. R., D. H. HABECK, AND D. L. MATTHEWS-Life History and Imma-
ture Stages of a Plume Moth Sphenarches anisodactylus (Lepidoptera:
Pterophoridae) in Florida ............................................................. 257
SCHUSTER, J., AND P. REYES-CASTILLo--Passalidae: New Larval Descriptions
From Taiwan, Philippine Islands, Brunei and Ivory Coast ................. 267
WIRTH, W. W., AND J. R. LINLEY-Description of Dasyhelea chani New Species
(Diptera: Ceratopogonidae) From Leaves of the Water Lettuce (Pistia
stratiotes) in Florida ..................................................................... 273
PETIT, F. L.-Distinguishing Larval Instars of the Vegetable Leafminer,
Liriomyza sativae (Diptera; Agromyzidae) ........................................ 280
PHILLIPS, T. W.-Responses of Hylastes salebrosus to Turpentine, Ethanol, and
Pheromones of Dendroctonus (Coleoptera: Scolytidae .......................... 286
of Feral Male Banded Cucumber Beetles to the Sex Pheromone 6,12-Di-
methylpentadecan-2-one ................................................................... 292
HALL, D. G.-Stand and Yield Losses in Sugarcane Caused by the Wireworm
Melanotus communis (Coleoptera: Elateridae) Infesting Plant Cane in
F lorida .................................................................................... 298
ATKINSON, T. H., P. G. KOEHLER, AND R. S. PATTERSON-Annotated Checklist
of the Cockroaches of Florida (Dictyoptera: Blattaria: Blattidae,
Polyphagidae, Blattellidae, Blaberidae) ......................................... 303

Continued on Back Cover

Published by The Florida Entomological Society

President .................................................................................. J. E Eger
President-E lect ........................................................................... J. F. Price
Vice-President ..................... ... ....................... J. L. Knapp
Secretary ...... .. ................... ................................... J. A. Coffelt
Treasurer ................................... ...................... .................... A. C. Knapp
Other Members of the Executive Committee
R. S. Patterson J. E. Pefia F. D. Bennett
M. Camara R. Coler
J. R. McLaughlin, USDA/ARS, Gainesville, FL ....................................... Editor
Associate Editors
Agricultural, Extension, & Regulatory Entomology
Ronald H. Cherry-Everglades Research & Education Center, Belle Glade, FL
Michael G. Waldvogel-North Carolina State University, Raleigh, NC
Stephen B. Bambara-North Carolina State University, Releigh, 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
John H. Brower-Stored Product Insects Research Laboratory, Savannah GA
Theodore E. Burk-Dept. of Biology, Creighton University, Omaha, NE
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
Omelio Sosa, Jr.-USDA Sugar Cane Laboratory, Canal Point, 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 M manager .................................................................... A. C. Knapp
FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and De-
cember. Subscription price to non-members is $30 per year in advance, $7.50 per copy;
institutional rate is $50 per year. Membership in the Florida Entomological Society,
including subscription to Florida Entomologist, is $25 per year for regular membership
and $10 per year for students.
Inquiries regarding membership and subscriptions should be addressed to the Busi-
ness Manager, P. O. Box 7326, Winter Haven, FL 33883-7326.
Florida Entomologist is entered as second class matter at the Post Office in DeLeon
Springs and in Winter Haven, FL.
Manuscripts from all areas of the discipline of entomology are accepted for consider-
ation. At least on. author must be a member of the Florida Entomological Society.
Please consult "Instructions to Authors" on the inside back cover.
This issue mailed June 29, 1990


The 73rd annual meeting of the Florida Entomological Society will be held August
5-9, 1990 at the Camino Real Hotel in Cancun, Mexico. Travel and hotel arrangements
are being handled through Holbrook Travel, 3540 N.W. 13th Street, Gainesville, FL
32609 (Phone 1-800-345-7111), Attn: Ms. Joyce Rickard. Registration forms and addi-
tional information will be mailed to members in the Newsletter.

For additional information contact:
Joseph L. Knapp, Chairman
Program Committee. FES
University of Florida
Citrus Research and Education Center
700 Experiment Station Road
Lake Alfred, FL 33850

A New Type of Article for our Authors and Subscribers

We are proud to announce that scientists may submit articles for publication in a
FORUM section of Florida Entomologist. FORUM articles (1-2 per issue) will appear
at the beginning of each issue in a section marked FORUM.
Articles for the FORUM section must follow the general style guidelines for all
other articles submitted to Florida Entomologist. FORUM articles must be of high
scientific quality, demonstrate acceptable experimental design and analysis, and cite
appropriate sources to support findings. FORUM articles will include "cutting edge"
science, scientifically meritorious but controversial subjects, new methodologies (de-
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documented challenges to existing entomological techniques, philosophies or experimen-
tal paradigms.
Submitted articles should include "Submitted to Florida Entomologist: FORUM"
on the title page. Three or more peer reviews will be acquired by the Associate Editor
for FORUM publications.
We feel the addition of a FORUM section will expand the scope of Florida En-
tomologist and allow readers and publishing scientists an additional creative outlet that
will complement our symposia, research articles, and notes.

Stansly & Sanchez: Biology & Behavior of Cydia fabivora 219


Southwest Florida Research and Education Center
P.O. Box 5127
Immokalae, FL 33934

Associaci6n de Productores de Ciclo Corto (APROCICO)
Casilla Postal 134
Quevedo, Ecuador


Cydiafabivora (Meyrick) is a neotropical pod and stem borer of common beans, lima
beans, and soybeans. All its life stages are larger than those of four similar pod boring
olethreutines which do not attack stems of their leguminous hosts. C. fabivora com-
pleted its life cycle in 29 days in the laboratory at ambient temperature on artificial diet
supplemented during the first larval stadium with fresh soybean seeds. Generation time
estimated from these results would allow three generations per soybean crop, first in
stems and then in seeds. Cage studies showed that before flowering, leaf undersides
were the predominant oviposition sites (55%), whereas pods were highly preferred
(84%) when available. Flexibility in the use of either stems or pods allowed the insect
to maintain itself throughout the crop cycle, increasing the risk to ripening seeds.


Cydia fabivora (Meyrick) es un barrenador neotropical de tallos y vainas del frijol
y la soya. Todos sus estadios de vida son mas grande en comparaci6n a cuatro barrena-
dores de vaina similares de la misma subfamilia que no atacan el tallo de sus huespedes
leguminosas. C. fabivora complete su ciclo biol6gico en 29 dias en el laboratorio a tem-
peraturas ambientales, alimentandose con una dieta artificial suplementada en el primer
estadio larval por granos tiernos de soya. Un tal desarrollo en el campo permitiria tres
generaciones por ciclo de soya, primero en tallos y luego en vainas. En ensayos de jaula
se observe que, antes de la floraci6n, se encontraron mas huevecillos (55%) en el env6s
de la hoja que en otra parte de la plant. Una vez disponibles, las vainas (84%) eran
preferidas para la oviposici6n. Flexibilidad al uso de tallos o vainas permiti6 al insecto
mantenerse durante todo el ciclo del cultivo aumentando el riesgo de dafio al grano.

Soybean production has increased in recent years on Ecuador's coastal plain, where
over 66,000 ha were planted in 1987 (del Salto & Tschirley 1988). Most of this production
is concentrated in the central region within a 50 km of radius of Quevedo, where climatic
and edaphic conditions allow 2 to 3 consecutive crops a year, each averaging ca. 2,000
kg/ha. Often, soybean at all stages of development can be found in close proximity. Such
intensive cropping practices may have contributed to the increased incidence of certain
pests in recent years, including Cydia fabivora (Meyrick). This species was first re-
ported attacking stems and pods of soybean on the coast of Ecuador by PAliz & Mendoza
(1980). C. fabivora is also known as a pest of soybean in other parts of tropical America
including Brazil, where it was initially confused with Epinotia aporema Walsingham
(Tortricidae: Olethreutinae) in many reports (Smith 1978). Forrester (1978) cited data

Florida Entomologist 73(2)

indicating the pest status of C. fabivora and the ineffectiveness of insecticidal control
against it.
Laspeyresiafabivora was described by Meyrick (1928) from a single male specimen
reared from common bean in Columbia. Synonyms of C. fabivora are L. leguminis
which Heinrich (1943) had described from a series of specimens collected in Peru,
Panama, and El Salvador (Clarke 1958), and Eulia prosecta Meyrick (Meyrick 1932,
Clarke 1972).
Although the species is widespread throughout Central and South America on beans,
lima beans, and soybeans (Clarke 1972), little further information exists in the literature
aside from brief mention in a review of olethreutines attacking grain legumes (Perrin
& Ezueh 1978). The objective of our study was to describe the insect's basic biology
including oviposition behavior, because this latter might bear on the eventual site of
larval feeding in either stems or pods.


All laboratory work was conducted at the Asociaci6n de Productores de Ciclo Corto
(APROCICO) in Quevedo, Los Rios Province. Field work was carried out at the
Pichilingue experiment station of the National Agriculture and Livestock Research
Institute (INIAP), 7 km SE of Quevedo (mean temperature = 24.3 C, mean RH =
83%). Temperature and humidity, monitored in the laboratory with a mechanical hydro-
thermograph, averaged 25 C (SD = 0.8, range = 240C to 27C) and 65% RH (SD -
19.3, range =42% to 96%).


A small laboratory colony was started from field-collected late instar larvae which
were placed in pairs in 59 ml clear plastic cups partly filled with sugarcane borer diet
(Bioserve Product "F" 9775). Reared pupae were placed in open petri dishes in rectan-
gular emergence/oviposition cages 35 cm high, by 45 cm square. The cages were framed
with lacquered wood, had hardware cloth tops and bottoms and a plywood side fitted
with a cloth sleeve and three glass sides. The insides were lined with wax paper for
oviposition. Eggs were cut from the lining, disinfected with a spray of 0.16% of benomyl
and drained on paper toweling for 5 min. Eggs were held on moistened filter paper
inside sterilized, parafilm-sealed glass petri dishes (105 x 15cm) that contained a mois-
tened cotton ball. Neonate larvae were fed fresh young soybean seeds that were re-
plenished every other day. Second instars were transferred with a camel hair brush to
59 ml plastic cups where they were maintained on sugarcane borer diet until pupation.
A pinto bean diet (Leppla 1985) was sometimes substituted for sugarcane borer diet in
routine colony maintenance. Otherwise, the same methods were employed for biological
studies as for routine rearing.


Head capsule width, egg length and egg width were measured with a stereoscopic
microscope fitted with an ocular micrometer. Head capsule size and the presence of
exuviae were noted every 12 h to determine the duration of each stadium for 20 individ-
uals which survived from egg to pupa. Larvae were weighed immediately after molting
with an electronic analytic balance accurate to 0.1 mg.
Pairs of laboratory-reared moths used to initiate the study were placed in square
450 ml wide-mouth glass jars (height = 15 cm, width = 10 cm) lined with wax paper
and closed with nylon netting fastened with a rubber band. Cotton balls soaked in a 3:1


June, 1990

Stansly & Sanchez: Biology & Behavior of Cydia fabivora 221

water to honey mixture provided a food source. Eggs were counted, measured and then
observed every 12 h to record color changes, viability, and incubation time. Neonate
larvae were maintained in groups of fives on fresh soybeans until the first molt and then
placed individually in diet cups. Pupae were sexed, measured (length and maximum
width), weighed and then placed in pairs on moistened filter paper in 450 ml oviposition
jars. Wingspan of adults was measured by spreading the wings on millimeter paper
under the microscope. Comparisons between male and female pupae and adults using
Student's T-test were made on the basis of 20 randomly chosen individuals of each sex.

Ovipositional Patterns

Six pots, each containing 2 soybean plants of the same age were placed in 1 m3 wood
frame cages covered with wire window screen. There were three cages, one for plants
in each of three development stages: vegetative (V8), flowering (R2), and pod-filling
(R6) (Fehr & Caviness 1977). Five moth pairs between 2 and 3 days old were released
into each cage and left for 48 h, after which the moths were removed and the plants
examined for eggs, noting the plant part on which oviposition had occurred.

Field Observations

Affected plant parts were dissected to determine the stage present and the site and
extent of damage. Field-collected eggs and larvae were reared in the laboratory as
previously described, and parasites collected.



Courtship and copulation took place ca. 48 h post-emergence; the pair assumed an
end to end position. Females began ovipositing almost immediately afterward, continu-
ing for 2 to 4 d (mean = 2.6 d, SE = 0.69). Eggs were glued to the substrate, either
singly or occasionally in small groups of 2 to 4. Mean egg production was 44 per female
(range = 32-56, SE = 1.5). The ventrally flattened eggs measured 0.89 mm in length
(range = 0.75-0.95, SE = 0.016) and 0.66 mm in width (range = 0.60-0.75, SE =
0.011), were pale yellow initially and covered with a raised hexagonal reticulation. Red
spots appeared below the chorion within 24 h of oviposition. These eventually coalesced
to make the entire egg red. Incubation period varied between 4 and 5 d (mean = 4.6
d, SE = 0.05), and viability was 82% (N = 100).
The larval integument was basically without pigment except for the prominent
prothorax and heart-shaped head, although neonate larvae were light orange in color.
Approximately 48 h of the last larval stadium was spent as a non-feeding prepupa.
Length, weight, development time, and head capsule width for the five larval instars
are given in Table 1.
The pupa had two conspicuous transverse bands of spines on abdominal sterna 3
through 9. Females were larger and heavier than males (Table 2). The pupal stage was
completed in 9.2 d regardless of sex (range = 8-11, SE = 0.24, t = 0.65). Thus the
average time from oviposition to adult emergence was 29.2 d (range = 25.5-32.5).
The adult moth has been adequately described elsewhere (Meyrick 1928, Heinrich
1943). Not surprisingly, females were again larger and heavier than males and also
survived longer (Table 3). If the midpoint of the adult female stadium (4 d) were as-
sumed to be the ovipositional midpoint, then the average generation time would be 33
d. Given 11 to 120 d from planting to harvest, there could be sufficient time for the
completion of three generations per soybean crop.

Florida Entomologist 73(2)


1 2 3 4 5

HEAD mean 0.34 0.74 1.05 1.4 1.9
CAPSULE min. 0.30 0.50 0.95 1.2 1.8
WIDTH max. 0.44 0.90 1.10 1.6 2.0
(mm) se 0.009 0.02 0.07 0.03 0.13

LARVAL mean 1.7 5.9 8.8 13.1 17.8
LENGTH min. 1.5 5.0 7.0 11.0 15.0
(mm) max. 1.9 6.8 10.0 14.5 20.0
se 0.29 0.14 0.2 0.20 0.27

LARVAL mean 1.0 3.1 10.7 50.4 80.8
WEIGHT min. 0.8 2.5 12.5 44.0 63.0
(mg) max. 1.2 3.5 8.5 60.0 94.5
se 0.02 0.054 0.22 0.98 1.95

DEVELOPMENT mean 3.9 2.2 2.8 2.4 4.1
TIME min. 3.3 2 2 2 3.5
(d) max. 4.5 3 3 3 4.5
se 0.09 0.07 0.06 0.07 0.06

Oviposition Behavior

Oviposition began after ca 48 h. Sixteen percent of the eggs were laid on stems of
plants at all three growth stages. In pre-flowering soybean, the remaining eggs were
laid predominantly on the undersides of leaves (55%), with leaf uppersides (16%) and
petioles (12%) accounting for the rest (Table 3). The same pattern occurred on flowering
plants except that flowers received 20% of the eggs at the expense of leaves. At pod-fill,
all 84% of eggs not on stems were found on pods. Thus, the preferred oviposition site
shifted from leaves to reproductive structures over the course of plant development.



PUPA 6 9 6 Y 5 Y
Length (mm) 9.3 10.1 10.5 11.0 6.9 8.9 5.1 0.001
Width (mm) 2.5 2.7 3.0 2.8 1.9 2.5 4.9 0.001
Weight (mg) 40.0 47.5 31.5 33.6 41.5 60.7 5.9 0.001
Length (mm) 7.6 8.8 7 8 8 10 10.1 0.001
Wingspread (mm) 15.8 17.8 17 19 15 19 10.8 0.001
Survivorship (d) 5.2 8.0 7 9 4 7 15.4 0.001

aStudents T-test used to determine statistical significance.

June, 1990

Stansly & Sanchez: Biology & Behavior of Cydia fabivora 223



PRE-FLOWERING 112 16% 55% 13% 16% -
FOWERING 93 12% 38% 13% 16% 22% -
POD FILL 104 16% 84%

Field Observations

First instar larvae attacking plants in vegetative stages began perforating the stem
soon after eclosion, often at the axil of the petiole, causing desiccation of the trifoliate.


C. C. Leguminovora C.
nigricana phychora glycinivorella fabivora

DISTRIBUTION: Europe Sub-sahara N.E. Asia Cen. & S.
N. America India America

HOST RANGE: peas common bean soybean common bean
lima bean lupin lima bean
pigeon pea soybean

PART ATTACKED seed seed seed stem and seed

length (mm) 0.75 0.45 0.48 0.89
width (mm) 0.53 0.35 0.35 0.66
incubation (d) 6-8 2-4 7-9 4-5
length (1) (mm) 1.2 0.7-0.9 0.8-1.1 1.73
length (5) (mm) 12-15 8.5-10.4 6-9.5 15-20
head (5) capsule
width (mm) 1.27 0.90 1.07 1.9
duration (d) 19-30 11-14 18-25 13-18

length (mm) 7-8 6 6-7 8-11
width (mm) 1.6-1.9 1.5 1.8 2-2.9
duration (d) 10-15 5-7 10-13 8-11

length (mm) 7-8 4-6 7 7-9
wingspan (mm) 11-13 14-16 13-15 15-19
survival (d) 10-12 4-7 10-13 4-9

aNumbers in parentheses indicate instar.

224 Florida Entomologist 73(2) June, 1990

Otherwise, the neonate larva penetrated the stem directly, leaving a short encircling
mine. The larva spun a silken support and remained in the same stem until development
was completed. Boring of the main stem killed small plants.
Attacked pods could be identified by short brownish mines where the first instar
larva had passed on its journey to the seed. Silken support webs were also spun in pods,
and one or two seeds were consumed during larval development, depending on seed
maturity. Some larvae were still feeding at harvest time. Pupae were normally found
in thin cocoons at the site of larval development in both stems and pods.
Hymenopteran parasitoids reared from C. fabivora included Trichogramma sp.
(Trichogammatidae) from eggs and Bracon sp., Apanteles sp., and Orgilus sp.
(Braconidae) from larvae.

Comparison with Related Species Attacking Legumes

C. fabivora is compared in Table 4 with three other olethreutine species that attack
grain legumes (Perrin and Ezueh, 1978). C. fabivora stands out by its relatively large
size and ability to use both stalks and pods. By feeding on either stems or seeds it could
complete three generations per crop cycle and potentially build up large populations
that would be difficult to control chemically because of the larva's cryptic habitat. Crop
rotation, or at least fallow periods between crops, and uniform planting dates to reduce
possible immigration from mature stands to new plantings, would probably provide
more effective and economical control.


Thanks are due the staff of APROCICO and the Pichilingue Experiment Station
(INIAP) for their cooperation, to D. H. Habeck for a helpful review and handling of
specimens for identification, and for species determinations to J. Heppner (C. fabivora)
and P. M. March (Braconidae), both of the U.S. National Museum, Washington, DC
20560. This work was supported by funds granted by the United States Agency for
International Development (USAID), to the Institute for Tropical Agriculture of the
University of Florida, contract No. 518-0032-C-00-1040. Florida Agricultural Station
Journal Series No. R-00565.


CLARKE, J. F. G. 1958. The correct name for a pest of legumes (Lepidoptera, Ole-
threutidae). Proc. Entomol. Soc. Washington., 60(4): 187.
1972. Two pests of beans from tropical America Lepidoptera: Olethreutidae).
Proc. Entomol. Soc. Washington., 74(4): 467-471.
DEL SALTO, R., AND D. TSCHIRLEY. 1988. Comercializaci6n en el sub-sector de
alimentos balanceados en el Ecuador, Vol. III: studio "La Soya a Nivel de
Finca". Documento T6cnico No 5. Institute de Estrategias Agropecuarios
(IDEA), Quito. 89 pp.
HEINRICH, C. 1943. A new species of Laspeyresia, a bean pest from tropical America
(Lepidoptera: Olethreutidae). Proc. Entomol. Soc. Washington, 45(3): 71-73.
FEHR, W. R., AND C. E. CAVINESS. 1977. Stages of soybean development. Iowa
Coop. Ext. Serv. Spec. Rep. 80: 12 p.
FORRESTER, L. A. 1978. Chemical control of soybean pests in Brazil, pp. 253-256 in
S. R. Singh, H. F. Van Emden, and T. A. Taylor [eds.], Pests of Grain Legumes:
Ecology and Control. Academic Press, NY.
LEPPLA, N. C. 1985. Anticarsia gemmatalis, pp. 189-196 in P. Singh and R. F.
Moore [eds.], Handbook of Insect Rearing, Vol. 2, Elsevier Science Puplishers
B. V., Amsterdam.

Howard: Natural Products Against Mahogany Webworm

MEYRICK, E. 1928. Exotic Microlepidoptera 3: 449.
1932. Exotic Microlepidoptera 4: 259.
PALIZ, V., AND J. MENDOZA. 1980. Informe annual t6cnico. INIAP, Estaci6n Tropical
Pichilingue, Departamento de Entomologia. P. 40.
PERRIN, R. M., AND M. I. EZUEH. 1978. The biology and control of grain legume
olethreutids (Tortricidae), pp. 201-218 in S. R. Singh, H. F. Van Emden, and T.
A. Taylor [eds.], Pests of Grain Legumes: Ecology and Control. Academic Press,
SMITH, J. G. 1978. Pests of soybean in Brazil, pp. 167-178 in S. R. Singh, H. F. Van
Emden, and T. A. Taylor [eds.], Pests of Grain Legumes: Ecology and Control.
Academic Press, NY.

-04w-pp - C L -c


University of Florida
Ft. Lauderdale Research & Education Center
3205 College Avenue
Ft. Lauderdale, Florida 33314


In laboratory tests, both Margosan-O, diluted to 20 ppm azadirachtin in water, and
Dipel 2X, which contains the entomopathogenic bacterium, Bacillus thuringiensis var.
kurstaki Berliner, at 9,600,000 international units (i.u.)/liter H20, applied to foliage of
West Indies mahogany, Swietenia mahagoni Jacquin, inhibited feeding by young
mahogany webworms, Macalla thyrsisalis Walker, as measured by differences in
growth and in the production of fecal pellets (P < 0.05). No clear effect was observed
on older larvae. In a field test, the mean number of larvae per tree was reduced 10-fold
(P < 0.05) ten days after treatments with either 20 ppm azadirachtin in H20 or B. t at
19,200,000 i.u./liter H20 applied to the foliage of West Indies mahogany. One or more
treatments with either of these materials during the spring when mahogany webworms
are active on foliage is a suitable method of controlling this pest.


En pruebas de laboratorio, Margosan-0, diluido a 20 ppm azadirachtin en agua, y
Dipel 2X, lo cual contiene el bacterium entomopatog6nico, Bacillus thuringiensis var.
kurstaki Berliner, a 9,600,000 unidades internacionales (u.i.)/litro H20, aplicado al fol-
laje de caoba antillana, Swietenia mahagoni Jacquin, inhibieron la alimentaci6n de lar-
vas de primeros estadios de Macalla thyrsisalis Walker, media por diferencias en
crecimiento y en la producci6n de pelotillas fecales (P < 0.05). No se observ6 un efecto
claro sobre las larvas mas crecidas. En un ensayo de campo, el promedio de larvas por
Arbol fue reducido diez veces (P < 0.05) diez dias despu6s de tratamientos con
azadirachtin a 20 ppm/litro HO, o B. t. a 19,200,000 u.i./litro H20 aplicado al foliage de
caoba antillana. Uno o mas tratamientos con cualquier de estos materials durante la
primavera cuando M. thyrsisalis es active sobre el follaje es un m6todo satisfactorio
para controlar esta plaga.


Florida Entomologist 73(2)

Larvae of the mahogany webworm, Macalla thyrsisalis Walker (Lepidoptera:
Pyralidae), attack foliage of mahoganies, Swietenia spp. (Meliaceae). In southern
Florida, where the West Indies mahogany, S. mahagoni Jacquin, is native and an
important ornamental, mahogany webworms usually occur on foliage beginning in April
of each year and become abundant during about five weeks when new leaves are expand-
ing. After June, larval populations become greatly reduced, apparently due at least
partly to the impact of parasitoids, including Apanteles sp., Habrobracon sp.
(Hymenoptera: Braconidae) and Lespesia sp. (Diptera: Tachinidae) (Howard & Solis
1989). The effects of defoliation on growth of mahoganies by mahogany webworms are
not known, but shrouding of the foliage in webbing and defoliation by the insect consti-
tute aesthetic damage to these ornamentals. Larval populations often become very
dense, and as the larvae mature, they drop onto cars, sidewalks, etc., causing a nui-
sance. Each year during this period, the Cooperative Extension Service receives numer-
ous requests for information on how to control this insect.
Environmentally compatible methods of suppressing larval populations of the
mahogany webworm are needed. Two natural products appeared to be good candidates
for this use.
Azadirachtin, and other compounds contained in the seed of the margosa, or neem
tree, Azadirachta indica A. Jussieu (Meliaceae), have antibiotic, insect antifeedant, and
insecticidal properties (Jacobson 1986, 1989, Warthen 1989, Pradhan & Jotwani 1968).
The neem tree is native to India and Burma and is planted as an ornamental or shade
tree in many tropical countries. Neem seed extracts have been shown to affect more
than 80 species of insects (Warthen 1989), but the toxicity of these substances to higher
animals is apparently extremely low. The oral LD, of azadirachtin for mice was re-
ported as 13,000 mg/kg (Jacobson 1989).
Entomopathogenic bacteria of the Bacillus thuringiensis group (B. t.) are specific
to lepidopterous and certain other larvae, and formulations using these bacteria are
recognized as among the safest and environmentally benign of natural pesticides (Coppel
& Mertins 1977).
The present paper reports results of laboratory tests and a field trial using
azadirachtin or B. t. to suppress mahogany webworm populations.


The laboratory test included four different treatments: (1) Margosan-O (W. R. Grace
& Co., Cambridge, Massachusetts) diluted in water to 20 ppm azadirachtin, (2) Dipel
2X (Abbott Laboratories, North Chicago, Illinois), which contains the HD-1 isolate of
B. thuringiensis var. kurstaki (Berliner) diluted with water to 9,600,000 international
units (i.u) per liter, (3) Plyac (Hopkins Agricultural Chemical Company, Madison, Wis-
consin 53707), which is a non-ionic spreader-sticker suitable for use with the above
materials, at the rate of 1 ml per liter H20, and (4) untreated controls. Plyac at 1 ml
per liter was added to the azadirachtin and B. t. mixtures to insure coverage of the
foliage. The B. t. was obtained from the manufacturers a few days prior to this study.
The azadirachtin was obtained 9 months earlier and stored in a laboratory at about 25
C. Materials were applied with a compressed air hand-held sprayer to young foliage of
new shoots selected at random on a West Indies mahogany tree. When almost dry,
sprayed foliage was clipped from the tree. One leaf was placed per Petri dish, with five
replications for each treatment. An early instar mahogany webworm 5-12 mm long was
placed on the leaf in each Petri dish. After three days, the number of fecal pellets was
determined for each larva as an indication of feeding activity. After one week, the
larvae were examined and measured. A similar experiment was conducted with nearly
fully grown larvae about 30 mm long.

June, 1990

Howard: Natural Products Against Mahogany Webworm 227

A field trial was conducted on West Indies mahoganies of about 3 to 4 m in height
at the Fort Lauderdale Research and Education Center. These were infested with
mahogany webworms which were mostly in early instars. The treatments were (1)
azadirachtin at the above rate, (2) B. t. at double the above rate, and (3) untreated
control. Plyac at 1 ml per liter was added to these mixtures. Treatments were assigned
at random to four trees per treatment. On May 10, 1989 and again 15 days later, the
materials were applied with a compressed air hand-held sprayer so as to fully cover the
foliage. Precipitation was recorded at a rain gauge about 200 m from the mahogany
trees. The numbers of webworms were determined just prior to spraying. The day after
application of the treatments, 30 medium to full-grown mahogany webworms were
placed on each tree to supplement the natural populations. The numbers of webworms
on each mahogany tree were determined two, ten, 15 and 20 days after spraying.
Results were analyzed with the Analysis of Variance and the Waller-Duncan Bayesian
k-ratio t-test (SAS Institute 1985).


No mortality was observed in treatments or control in the laboratory test with
nearly full-grown larvae. There were no significant differences in the numbers of fecal
pellets produced by larvae on foliage treated with azadirachtin or B. t., compared to
the controls. These larvae increased only slightly in length and pupated within 7 days
of the commencement of the experiment.
There were significantly fewer fecal pellets produced by younger larvae over a 3
day period on foliage treated with B. t. and with azadirachtin, compared with the
controls (P < 0.05). The mean increase in length of mahogany webworms on foliage
treated with either azadirachtin or B. t. was less than 1/4 of that of webworms in the
controls (P < 0.05) (Table 1). These results indicate that both B. t. and azadirachtin
curtailed feeding of younger mahogany webworms. There was no significant difference
in mean numbers of pellets or mean increase in length between the spreader-sticker
treatment and the control.
Three days after the application of the materials there was no mortality of young
larvae in any treatment including the control. At the end of seven days, three of the
five larvae on foliage treated with azadirachtin and one on foliage treated with spreader-
sticker were dead. All larvae in the other treatments were alive. In an earlier study,
Reinert & Howard (1982) observed that mahogany webworms were alive after 24 hours
exposure to B. t., but that study did not reveal that this treatment curtailed feeding.


Mean number of Mean growth of
fecal pellets SD larvae
Treatment (3 days)a SD (7 days)b SD

Control 144.3a 26.9 11.6a 2.2
Spreader-sticker 114.7ab 11.2 8.4a 4.8
Azadirachtin 84.8bc 35.9 1.8b 2.0
B. t. 62.4c 14.2 3.2b 2.6

a'bMeans within the column not followed by the same letter are significantly different [" F 10.66, df 16, P
< 0.05, k ratio = 100; b F = 8.68, df = 16, P < 0.05, k ratio = 100 ( Waller-Duncan Bayesian k-ratio t-test, SAS
Institute 1985)].

228 Florida Entomologist 73(2) June, 1990

In the field test, on the second day after application of the materials to West Indies
mahogany trees, populations of this insect had increased about equally in all treatments
including controls, probably partly due to the placement of 30 medium to full-grown
larvae per tree, but also due to natural infestation as evidenced by an apparent increase
in the numbers of early instar larvae. Ten days after the first treatment application,
the populations of mahogany webworms per tree in both azadirachtin and B. t. treat-
ment groups were reduced ten-fold or more compared to the control group (P < 0.05,
F = 9.50, df = 9). Mean webworms per tree by treatments were as follows:
azadirachtin, 2.3 (SD = 3.86); B. t., 3.0 (SD = 1.14); and control, 34.3 (SD = 20.1)
(Fig. 1). Larvae on treated and untreated trees were of mixed instars. Since the larval
stage of mahogany webworms lasts about ten days in the field (Howard & Solis 1989),
it is unlikely that later instar larvae placed on the trees the day after the application of
treatments were still present in the population ten days after treatment applications.
At this point, the webworm populations presumably included full-grown webworms
that were in early instars and naturally established on the trees before spraying and
younger webworms that became established naturally since the application of treat-
ments. Since the laboratory tests showed that the treatments curtail feeding and are
most effective against younger larvae, it is presumed that most of the younger larvae
attempting to feed on foliage treated with either azadirachtin or B. t. probably dropped
off after a few days. There was noticeably more webbing and feeding damage on un-
treated trees than on trees treated with either material. Populations in both
azadirachtin and B. t. treatment groups remained low 15 days after the initiation of
treatments. Populations had also declined in the control group, but since some early
instar larvae were observed in all treatment groups, and both materials are generally
not persistent under field conditions, the treatments were repeated at this time to
prevent re-establishment of webworm populations. On 31 May, twenty days after the
first treatment application, mahogany webworm populations on trees in the control
group had diminished as predicted from studies of their population dynamics (Howard

35 A Azadirachtin
S40 Bacillus thuringiensis
S0 Control
5 25-

0 20-

S 15.
z 10-
|2nd application
0 1 st application
1 5

0 2 10 15 20

Fig. 1. Mean numbers of mahogany webworms per West Indies mahogany tree
treated with azadirachtin, Bacillus thuringiensis, and control, Fort Lauderdale,
Florida, May 1989.

Howard: Natural Products Against Mahogany Webworm 229

& Solis 1989) and were about equal to those on trees treated with either of the two
The azadirachtin and B. t. treatments were effective in spite of rainfalls that oc-
curred during the field test. There was a heavy rainfall (40.1 mm) five days after the
first treatment applications. In addition there were seven days each with a mean of 0.35
mm of rainfall during the period of the test. Based on the results of this study, both
azadirachtin and B. t. are promising materials for the management of mahogany web-
worms. One or more applications in the spring "webworm" season would be adequate
for controlling this pest in Florida. The first application should be made at the beginning
of the leaf flushing period of mahogany when mahogany webworms are first observed
on foliage. In Florida, mahogany webworms are ojectionable to the public when they
occur in dense populations or cause aesthetic damage to mahogany trees. Thus, the need
for additional applications should be based on a subjective analysis of field observations.
Both materials, because of their low environmental persistence and low level of toxicity
to higher animals (Jacobson 1989) are highly suitable for protecting mahoganies in urban
environments. These products would also have advantages for use on timber trees.
However, although the mahogany webworm is widely distributed on mahoganies in
Tropical America (Howard & Solis 1989), it has thus far not been reported as a pest of
mahoganies under forest conditions.


I thank Mr. J. V. DeFilippis for technical assistance and the following for useful
technical information concerning their products: Mr. Robert O. Larson (Vikwood, Ltd.,
the developers of Margosan-O), Dr. James F. Walter (W. R. Grace & Co.) and Dr.
Joseph P. O'Connor (Abbott Laboratories). Drs. Monica Elliott and Nan Yao Su (Uni-
versity of Florida) kindly reviewed the manuscript. Mention of a product does not
constitute a guarantee by the University of Florida, nor imply endorsement of its use
in lieu of other similar products. This is Florida Agricultural Experiment Station Journal
Series No. 10056.


COPPEL, H. C., AND J. W. MERTINS. 1977. Biological insect pest suppression.
Springer-Verlag, Berlin, Heidelberg, New York.
HOWARD, F. W., AND M. A. SOLIS. 1989. On the distribution, life history, and host
plant relationships of the mahogany webworm, Macalla thyrsisalis Walker
(Lepidoptera: Pyralidae). Florida Entomol. 72: 469-479.
JACOBSON, M. 1986. The neem tree: natural resistance par excellence, pp. 220-232,
in M. B. Green and P. A. Hedin (eds.), Natural Resistance of Plants to Pests;
Roles of Allelochemicals. ACS Symposium series 296, American Chem. Soc.,
Washington, D.C.
JACOBSON, M. (ed.) 1989. Focus on phytochemical pesticides. Vol. I. The Neem Tree.
CRC Press.
KUBO, I., AND J. A. KOLCKE. 1982. Isolation of phytoecdysones as insect ecdysis
inhibitors and feeding deterrents, pp. 329-3246, in P. A. Hedin (ed.), Plant Re-
sistance to Insects. ACS Symposium Series 208, American Chem. Soc.,
Washington, D.C.
PRADHANS, S., AND M. G. JOTWANI. 1968. Neem as an insect deterrent. Chem. Age
India 19: 756-760.
REINERT, J. A., AND F. W. HOWARD. 1982. Susceptibility of mahogany webworm
to insecticides. Proc. Florida State Hortic. Soc. 95: 288-289.
SAS INSTITUTE. 1985. SAS user's guide: statistics. SAS Institute, Cary, N.C.
WARTHEN, J. D. 1989. Neem (Azadirachta indica A. Juss.): organisms affected and
reference list update. Proc. Entomol. Soc. Washington 91: 367-388.

Florida Entomologist 73(2)


University of Florida
Institute of Food and Agricultural Sciences
Citrus Research and Education Center
700 Experiment Station Road
Lake Alfred, FL 33850 USA


Copper hydroxide (Kocide 101, 50% metallic copper) did not affect the residual effi-
cacy of the acaricides, AGRI-MEK, Vendex, and A-1335, used to control citrus rust
mite. All acaricides, with or without copper, gave significant knockdown of rust mite
populations within 10 days after application, and their residual control exceeded 10
weeks. No compatibility problems were detected between acaricides and copper hy-
droxide. Copper sprays alone did not incite an increase in citrus rust mite populations
in all three experiments. The pretreatment population densities of citrus rust mite
differed among experiments but had no effect on the residual control of the acaricides.


Hydr6xido de cobre (Kocide 101, 50% cobre metalico) no afect6 la eficacia residual
de los acaricidas AGR-MEK, vendex, y de A-1335, que se usan para controlar el Acaro
de la rolla citrica. Todos los acaricidas, con o sin cobre, tuvieron un efecto rapido y
fulminante en la poblaci6n de Acaros dentro de los primeros 10 dias de su aplicaci6n, y
su control residual se extendi6 10 semanas. No se detectaron problems de com-
patibilidad entire los acaricidas y el hidr6xido de cobre. El rocio de cobre solo no incit6
un aumento en la poblaci6n de acaros en los tres experiments. La densidad de la
poblaci6n de Acaros antes del tratamiento fu6 distinta entire los experiments pero no
tuvo efecto en el control residual de los acaricidas.

Citrus rust mite, Phyllocoptruta oleivora (Ashmead) (Acari: Eriophyidae), is the
most serious mite pest of citrus in Florida (Yothers & Mason 1930, McCoy & Albrigo
1975, McCoy et al. 1988). Its feeding on a leaf can cause localized degreening of the
upper surface, degreened necrotic spots on the lower surface, and subsequent defolia-
tion (McCoy 1976). Fruit damaged by the extensive feeding of cirus rust mite develop
severe blemishes and discoloration (russet) on surface. Severe injury can result in smal-
ler fruit, lower juice volume, alteration in soluble solids, and premature fruit drop,
thereby reducing yield and fresh market value (McCoy et al. 1976b, Allen 1976). Since
the increase of citrus rust mite populations in spring and summer coincide with that of
important citrus fungal diseases such as melanose and greasy spot (Whiteside 1988),
acaricides are usually mixed with copper fungicides. Consequently, the efficacy of differ-
ent acaricides may be influenced by the addition of copper, or citrus rust mite population
dynamics may be affected by the frequent application of copper fungicides alone
(Johnson 1960a, McCoy 1977, Dean 1979).
Winston et al. (1923) first reported that citrus rust mite was more abundant on
copper sprayed citrus than on unsprayed citrus. Griffiths & Fisher (1949) indicated that
zinc sulfate provided less control of citrus rust mite when mixed with lime. A mixture


June, 1990

Lye et al.: Copper Affects Citrus Rust Mite Control

of zineb (zinc ethylene-bis-dithio-carbamate) and copper resulted in less control of citrus
rust mite than zineb alone (Johnson et al. 1957). In studying the compatibility of several
fungicides with copper, Johnson (1960b) concluded that copper sprays prolonged high
population of citrus rust mite or caused a continuous increase when applied at the time
unsprayed populations were declining. In Texas, Dean (1979) reported that residual
control of citrus rust mite was reduced when copper (77% WP Kocide 101, cupric hydro-
xide) or 24.5% EC Supracide was added to other acaricides. However, McCoy et al.
(1989) in a study of the effects of spray volume and acaricide rate on the control of citrus
rust mite reported that the mean cumulative mite days on leaves treated with copper
(tracer) were significantly lower than on untreated leaves.
Since the chemicals and formulations of acaricides have changed during the past two
decades, it is appropriate to investigate the effects of copper on the efficacy of the
newly-developed miticides for the control of citrus rust mite. The objectives of this
study were to (1) evaluate the influence of copper on the residual efficacy of several
acaricides at defined pretreatment mite densities and (2) investigate the possible incita-
tion of copper spray alone to the increase of citrus rust mite populations on grapefruit
and 'Sunburst' mandarin. 'Sunburst' mandarin was recently reported to be more suscep-
tible to citrus rust mite, Texas citrus mite (Eutetranychus banksi (McGregor)), black
scale (Sassetia neglecta (Delotto)), and citrus mealy bug (Planococcus citri (Risso)) than
most cultivars (Albrigo et al. 1987). Therefore, the application of copper fungicides and
acaricides on this cultivar for the control of fruit diseases and citrus rust mite requires
more attention.

In the first and third tests, treatments were AGRI-MEK 0.15EC (mixed with 0.25%
FC 435-66 oil at 5.67 g ai/acre) at 31.5 ml per 378 liters water, Vendex 4L (566.99 g
ai/acre) at 35.5 ml per 378 liters water, untreated check, and combinations of the three
each with Kocide 101 (copper hydroxide, 50% metallic copper, 3.63 kg ai/acre) at 770 g
per 378 liters of water. In the second test, treatments were all the possible combinations
of four rates (15.7, 22.8, 30.2, or 45.5 ml per 378 liters water) of 'A-1335 25L' (an IGR
developed by Uniroyal Chemical Company, Inc) and untreated check with or without
Kocide 101 (3.63 kg ai/acre at 770 g per 378 liters water).
The first test was conducted in a 'Sunburst' mandarin grove in Avon Park, FL from
May through August 1988. 'Sunburst' trees were 5-yr-old and were ca. 2.43 m in height.
The test was conducted with a randomized complete block design with two trees per
treatment in each of six blocks. Ten fruit per tree were randomly selected from outside
tree canopy at or below ca. 160 cm height for counting citrus rust mite at one day before
treatment and at 10, 20, 30, 40 50 and 60 days posttreatment. The second and third
tests were conducted in two different 'Marsh' grapefruit groves in Davenport, FL from
June through August and July through September 1988, respectively. Grapefruit trees
in both tests were over 30-yr-old with rough lemon root stock. Both tests were con-
ducted with a randomized complete block design with one tree per treatment in each
of four blocks. In all three tests, treatments were applied with a spray volume of 1,890
liters water/acre on foliage until run off with a truck-mounted handgun sprayer at 400
psi. Twenty-five fruit per tree were randomly selected from outside tree canopy at or
below ca. 160 cm high for mite count at 1 week pretreatment and at 1, 2, 4, 6, 8, and
10 weeks posttreatment. Citrus rust mite populations on selected fruit were monitored
in the field by counting individual mites within two randomly selected 1-cm2 lensfield
areas on the shaded site of fruit with a 10X hand lens.
Data in Tests 1 and 3 were subjected to the /x+1 transformation before the
analysis of variance (ANOVA) was applied; however, means presented in Tables 1 and
3 for Tests 1 and 3, respectively, are means before the transformation. In each test, an
ANOVA was first applied on factorial treatments to evaluate the acaricide and copper

Florida Entomologist 73(2)

effects and another ANOVA was applied on the combined acaricide and copper treat-
ments (SAS Institute, 1988). Pretreatment data were used as sources of covariance in
the ANOVA to evaluate the treatment effects within each sampling date, and in the
repeated measures analysis (SAS Institute, 1988) to evaluate the overall residual treat-
ment effects through time. Waller-Duncan K-ratio t test was applied in Tests 1 and 3
to compare means among treatments (SAS Institute, 1988). Relationship between treat-
ment means and different rates of A-1335 in Test 2 was quantified by polynomial
contrasts in order to reveal the responding trend of citrus rust mite control as the rates
of A-1335 changed.


All pretreatment mite counts within each test were not significantly different among
combined acaricide and copper treatments. Block effect at pretreatment was significant
(F = 7.22, df = (4, 17), P < 0.01) in Test 1, but was not significant in Tests 2 and 3.
The pretreatment covariance and interaction of acaricide and copper were not significant
on each sampling date in each test.
The addition of copper hydroxide did not affect the efficacy of AGRI-MEK 0.15EC
and Vendex 4L in the control of citrus rust mite in the first test (Table 1.). All mite
populations in acaricide-treated plots were significantly lower than those in check plots.
Citrus rust mite populations of both checks (with or without copper) were significantly
different at 20, 30 and 40 days posttreatment, though both populations exhibited a
similar trend of increase until 60 days posttreatment.
All A-1335 treatment effects in the second test were significant (Table 2). The addi-
tion of copper hydroxide showed no effect to the four rates of A-1335 in the control of
mite populations. Population dynamics of citrus rust mite on both copper-treated and
untreated checks were similar. Mite populations in the copper-treated checks appeared
to be lower than those in the copper-untreated checks before 8 weeks posttreatment,
but the differences were not significant across the test period.
Relationships between acaricide rates and mean mite densities on fruit were signif-
icantly linear (F = 101.43, df = (1, 26), P < 0.01, contrast comparison) with regard to
an overall significant time effect (Wilk's lamda = 0.46, F = 5.20, df = (5, 22), P <
0.01) on the residual efficacy of A-1335. All A-1335 treatments, with or without copper,
significantly (F = 17.58, df = (1, 26), P < 0.01, compared with checks) controlled the
citrus rust mite populations to a very low level through time. Citrus rust mite popula-
tions in the 15.7 ml/acre treatment resurged slightly after 2 weeks posttreatment but
started to decline 4 weeks later whereas mite populations in the 45.5 ml/acre treatment
remained at a very low level after 2 weeks posttreatment, indicating a longer residual
activity of A-1335 applied at higher rates than at lower rates.
In the third test, effects of AGRI-MEK 0.15EC and Vendex 4L, with or without
copper, were also significant except at 4 weeks posttreatment (Table 3). This result was
consistent with the first test in which the addition of copper did not affect the efficacy
of these two acaricides. Mite populations on the copper-treated and untreated checks
fluctuated with a similar trend except that after 8 weeks posttreatment mite populations
on the copper-treated checks declined significantly whereas mite populations on the
untreated checks continued to increase. Causes of this rapid population declination on
copper-treated check trees were unknown; however, the differences in mite density
between copper-treated and untreated checks were not significant across the test period
except at week 12.
Citrus rust mite populations on the acaricide-treated trees (with or without copper)
remained nearly zero during posttreatment period and did not start to resurge until 50
days or 8 weeks posttreatment in all three tests (Tables 1, 2, and 3). These results


June, 1990

Lye et al.: Copper Affects Citrus Rust Mite Control

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Florida Entomologist 73(2)


Mean mite per cm2 surface on fruit
Rate Weeks after treatment
(formulated Copper' Pre-
ml/acre) addition treatment 1 2 4 6 8 10

15.7 No 7.60 4.64 0.53 2.40 6.00 4.14 1.89
15.7 Yes 4.58 5.91 0.50 2.87 8.21 5.78 2.15
22.8 No 4.62 6.09 0.59 1.31 2.78 2.24 1.56
22.8 Yes 8.70 5.44 0.40 2.38 5.68 2.46 1.15
30.2 No 4.87 3.59 0.13 1.14 2.64 1.08 0.70
30.2 Yes 8.96 3.78 0.61 2.11 1.99 3.34 2.10
45.5 No 9.64 5.14 0.64 0.55 0.78 0.54 0.52
45.5 Yes 8.72 1.56 0.10 0.27 0.49 0.92 0.91
Check No 11.74 20.00 22.97 52.20 63.76 18.46 4.00
Check Yes 8.78 11.00 15.79 32.14 42.93 24.40 5.35

ANOVA2: Acaricide F = 4.69 16.74 20.52 40.35 8.52 9.71
P>F <0.01 <0.01 <0.01 <0.01 <0.01 <0.01
Copper F = 2.26 0.91 1.15 1.21 0.65 1.79
P>F 0.14 0.35 0.29 0.28 0.43 0.19

'Kocide 101 (copper hydroxide, 50% metallic copper) at 3.63 kg ai/acre.
2Degree of freedoms for acaricide effect are 4 and 26; for copper effect are 1 and 26.

demonstrated that an effective acaricide, if applied correctly at an appropriate time,
will prevent the citrus rust mite populations from building up in summer.


Copper effects on the residual efficacy of AGRI-MEK 0.15EC, Vendex 4L and A-
1335 could not be detected in this study because copper-incorporated or regular
acaricides had the same high level of citrus rust mite control. The knockdown effects
of acaricides were also significant regardless of the addition of copper hydroxide. These
results revealed there were no detrimental chemical interactions between the acaricides
and copper hydroxide used in this study. McBride (1958) reported that the addition of
copper sulfate or copper oxide to zineb used for citrus rust mite control resulted in a
chemical reaction that reduced the zinc compound deposited by the spray. This study
has demonstrated that the tested acaricides gave sufficient citrus rust mite control for
at least 10 weeks or 60 days. Others & Mason (1930) suggested controlling the citrus
rust mite in early June and late September if a resurgence of the mite population
Citrus rust mite population dynamics on the untreated citrus were different among
these three experiments; however, they exhibited a similar trend, that is, populations
increased in midsummer and started to decrease in August as had been reported by
Others & Mason (1930), Pratt (1957), and McCoy et al. (1976a). With only a few
exceptions, mite population dynamics on the copper-treated checks coincided with those
on copper-untreated checks and were not significantly different in all three tests across
the test period. The copper hydroxide spray evidently did not incite the increase of mite
populations on acaricide-untreated checks in this study. On the contrary, mite densities
on copper-treated check were lower than on copper-untreated check in most of the
sampling dates in Tables 1 and 2, indicating that copper spray may have detrimental


June, 1990

Lye et al.: Copper Affects Citrus Rust Mite Control

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Florida Entomologist 73(2)

effect on the citrus rust mite populations. Since copper was applied early when the mite
population started to increase, our results cannot be compared with the results obtained
by Johnson (1960b) where a copper spray prolonged high densities of citrus rust mite
when applied to a declining population. Nevertheless, his suggestion that copper sprays
may physiologically affect the plants resulting in an increase of citrus rust mite popula-
tion cannot be applied to our results of no copper effects on the citrus rust mite popula-
Copper fungicide may incite the citrus rust mite population to increase on citrus
when the natural enemy of citrus rust mite, Hirsutella thompsonii, is suppressed by
the copper (Yothers & Mason 1930, Griffiths & Fisher 1949). Van Brussel (1975) also
reported that the increase in incidence of Hirsutella-infected citrus rust mite coincides
with the decrease in mite count on citrus in Surinam. McCoy et al. (1976a) reported
that an epizootic of this pathogen appeared to be responsible for a sharp decline in
populations of citrus rust mite during the summers of 1972 and 1973. Furthermore, the
abundance of H. thompsonii is density dependent to the citrus rust mite (Muma 1955,
McCoy 1981). The declination of citrus rust mite populations on most of the acaricide-un-
treated checks and the slow resurgence of citrus rust mite populations on acaricide-
treated citrus after ca. 6 weeks posttreatment in this study might be due to the increase
of this natural enemy in citrus groves thereafter. A further study is needed to evaluate
the effect of copper on the population of H. thompsonii and to quantify its relationship
with the population of citrus rust mite in the field.


The senior author appreciates the Merck & Co., Inc., for the support of his postdoc-
toral fellowship on the research of citrus rust mite.
Florida Agricultural Experiment Station Journal Series No. R-00366.


ALBRIGO, L. G., C. W. MCCOY, AND D. P. H. TUCKER. 1987. Observations of
cultivar problems with the 'Sunburst' mandarin. Proc. Florida State Hort. Soc.
100: 115-118.
ALLEN, J. C. 1976. A model for predicting citrus rust mite damage on Valencia orange
fruit. Environ. Entomol. 5: 1083-1088.
DEAN, H. A. 1979. Citrus rust mite control affected by certain pesticides. J. Rio
Grande Valley Hort. Soc. 33: 55-62.
GRIFFITHS, J. T., JR., AND F. E. FISHER. 1949. Residues on citrus trees in Florida.
J. Econ. Entomol. 42: 829-833.
JOHNSON, R. B. 1960a. The effect of copper compounds on the control of citrus rust
mite with zineb. J. Econ. Entomol. 53: 395-397.
JOHNSON, R. B. 1960b. The effects of copper on rust mite control with four rust mite
miticides. Proc. Florida State Hort. Soc. 73: 84-89.
JOHNSON, R. B., J. R. KING, AND J. J. MCBRIDE, JR. 1957. Zineb controls citrus
rust mite. Proc. Florida State Hort. Soc. 70: 38-48.
MCBRIDE, J. J., JR. 1958. Reaction of zineb with copper compounds, oil deposits
when applied with zineb, and deposits of zineb when applied with a variety of
materials. Proc. Florida State Hort. Soc. 71: 118-122.
McCoY, C. W. 1976. Leaf injury and defoliation caused by the citrus rust mite,
Phyllocoptruta oleivora. Florida Entomol. 59: 403-410.
McCOY, C. W. 1977. Horticultural practices affecting pytophagous mite populations
on citrus. Proc. Int. Soc. Citriculture 2: 459-462.
McCOY, C. W. 1981. Pest control by the fungus Hirsutella thompsonii, p. 449-512 in
H. D. Burges [ed.], Microbial control of pests and plants diseases 1970-1980.
Academic Press, New York.


June, 1990

Lye et al.: Copper Affects Citrus Rust Mite Control

McCoY, C. W., AND L. G. ALBRIGO. 1975. Feeding injury to the orange caused by
the citrus rust mite, Phyllocoptruta oleivora (Prostigmata: Eriophyoidea). Ann.
Entomol. Soc. Amer. 68: 289-297.
McCoY, C. W., L. G. ALBRIGO, AND J. C. ALLEN. 1988. The biology of citrus rust
mite and its effects on fruit quality. The Citrus Industry. Sept. 1988, p. 44-54.
McCoY, C. W., R. F. BROOKS, J. C. ALLEN, AND A. G. SELHIME. 1976a. Manage-
ment of arthropod pests and plant diseases in citrus agroecosystems. Proc. Tall
Timbers Conf. on Ecol. Animal Control by Habitat Manage. 6: 1-17.
McCoY, C. W., P. L. DAVIS, AND K. A. MUNROE. 1976b. Effects of late season fruit
injury by the citrus rust mite, Phyllocoptruta oleivora (Prostigmata:
Eriophyoidea), on the internal quality of Valencia orange. Florida Entomol. 59:
McCoy, C. W., B. H. LYE, AND M. SALYANI. 1989. Spray volume and acaricide rate
effects on the control of the citrus rust mite. Proc. Florida State Hort. Soc.
102:(In press).
MUMA, M. H. 1955. Factors contributing to the natural control of citrus insects and
mites in Florida. J. Econ. Entomol. 48: 432-438.
PRATT, R. M. 1957. Relation between moisture conditions and rust mite infestations.
Proc. Florida State Hort. Soc. 70: 48-51.
SAS INSTITUTE. 1988. SAS user's guide: Statistics. SAS Institute, North Carolina.
VAN BRUSSEL, E. W. 1975. Interrelations between citrus rust mite, Hirsutella
thompsonii and greasy spot on citrus in Surinam. Agric. Exp. Sta. Surinam Bull.
No. 98.
WHITESIDE, J. 0. 1988. Grove practices to prevent rind blemishes caused by fungual
diseases. Inst. of Food and Agric. Sci., Univ. of Florida, Florida Coop. Ext.
Serv. Packinghouse Newsletter No. 155, Gainesville, Florida.
WINSTON, J. R., J. J. BOWMAN, AND W. W. YOTHERS. 1923. Bordeaux-oil emulsion.
U.S. Dept. Agric. Bull. No. 1178. p. 1-24.
YOTHERS, W. W., AND A. C. MASON. 1930. The citrus rust mite and its control. U.S.
Dep. Agric. Tech. Bull. No. 176. p. 7-16.


University of Florida, IFAS
Citrus Research and Education Center
700 Experiment Station Road
Lake Alfred, Florida 33850

Florida Department of Citrus
Citrus Research and Education Center
700 Experiment Station Road
Lake Alfred, Florida 33850


Gamma radiation effects on Fuller rose beetle, Pantomorus cervinus (Boh.), eggs
were evaluated. Two age ranges of 1-5 and 6-12 day-old eggs were subjected to irradi-


Florida Entomologist 73(2)

ation doses of 0.075, 0.150, and 0.300 kilo Gray (kGy). Three trials were replicated with
a minimum of 50 eggs per dose per age per trial. One hundred percent mortality of all
1-5 day-old eggs was effected by all three irradiation doses. No egg hatch was observed
for the 6-12 day-old eggs irradiated with 0.30 kGy. A statistically insignificant percent
of hatch of 6-12 day-old eggs irradiated with 0.075 and 0.150 kGy occurred (16.3 and
0.10%, respectively). Dose was highly correlated to mortality.


Se evaluaron los efectos de radiacion gama sobre huevos del escarabajo de la rosa,
Pantomorus cervinus (Boh.). Huevos de 1-5 y de 6-12 dias de edad se sometieron a
d6sis de radiaciones de 0.075, 0.150, y a 0.300 kilo Gray (kGy). Se replicaron tres
ensayos con un minimo de 50 huevos por dosis por edad por ensayo. Un 100% de
mortandad de todos los huevos de 1-5 dias se obtuvo por las tres dosis de radiaciones.
No se observ6 eclosi6n de ninglin huevo de 6-12 dias de edad radiados con 0.30 kGy.
Ocurri6 un porciento estadisticamente insignificant de eclosi6n de huevos de 6-12 dias
de edad radiados con 0.075 y 0.150 kGy (16.3 y 0.10% respectivamente). La dosis estuvo
altamente correlacionada a la mortandad.

The Fuller rose beetle, Pantomorus cervinus (Boheman), is widely distributed
throughout the world and has a wide range of hosts including citrus (Anonymous 1966).
The beetle has a propensity for laying the majority of its eggs under the calyx of citrus
fruit (Coats & McCoy 1989), which has created a major problem for the U.S. citrus
industry. Japan has refused to accept any infested fruit, establishing a zero tolerance
for viable rose beetle eggs on fresh fruit imported into their country by 1990. This policy
has necessitated the implementation of reliable, highly effective, quarantine treatments.
Fumigation by the Japanese at the port of entry with methyl bromide (MeBr) has
resulted in large monetary losses to the U.S. citrus industry. Not only is fumigation
costly, but MeBr phytotoxicity causes rapid fruit deterioration (Haney et al. 1987,
Anonymous 1988). Furthermore, the use of chemical pesticides is not favorably received
by the consuming public. The U.S. Environmental Protection Agency banned ethylene
dibromide in 1987 and is requiring the re-registration of MeBr. There is currently no
quarantine treatment available to the U.S. citrus industry against Fuller rose beetle.
Numerous studies have shown radiation to be a reliable quarantine treatment for
control of insects in fruits and vegetables. Koidsumi (1930) first determined x-rays could
be successfully applied to effect mortality of insects in fruits in quarantine. Balock &
Christenson (1956) found both x-rays and gamma radiation to be effective in killing
fruit-infesting insects. Subsequent research detailed the dosages required to effect mor-
tality of specific insects (Burditt 1982, Burditt et al. 1981, Couey 1983, Kader et al.
1984, McMullen & Yeager 1982). A range of 0.05-0.75 kilo Gray (kGy) (1 kGy = 100
kRad) was determined as the effective insect-disinfestation doses for almost all insects
(Kader 1986).
Major radiation costs are due to the transportation of the products to and from the
radiation facility, and the cost of irradiation itself is very dependent on the throughput.
Costs of a cobalt-60 facility operated for 8000 h/yr can be as low as 5 cents/kg of product
per 10 kGy dose and 1,000,000 tons of output, or as high as $3.50/kg for the same dose
and only 2,000 tons of output (Brynjolfsson 1989). The feasibility of radiation is proven
by the fact that 38 operational facilities already exist in 24 countries, including four in
the U.S. The most extensive operation was established in Shihoro, Japan in 1973 and
currently has a monthly capacity to process 10,000 tons of potatoes (Matsuyama &
Umeda 1983).

June, 1990


Coats & Ismail: Ovicidal Effects of Gamma Radiation

Because gamma irradiation has been proven to be an effective, safe, and potentially
economical quarantine treatment for fruit-infesting insects, the present study was de-
signed to evaluate gamma radiation as an ovicidal treatment for Fuller rose beetle eggs.


Three separate radiation trials were completed. Because variable sensitivity of eggs
has been demonstrated, 1 to 5 and 6 to 12 days old were used. These age ranges were
chosen so the pre-hatch period of the Fuller rose beetle eggs could be evaluated. (Egg
hatch generally begins on day 15 at 280C and a buffer for earlier hatch at 13 and 14 days
was allowed.) Also, because variable sensitivity of eggs has been demonstrated, at least
two age ranges were necessary to evaluate this sensitivity in the Fuller rose beetle.
Three replications each consisting of a minimum of 50 eggs per dose per trial were
used with each age. Eggs laid by adults on wax paper strips were collected, counted,
and placed on a 20 mil screen mesh. This screen mesh was suspended above 1 cm of
distilled water in presterilized 7 dram plastic vials. The vials were capped and irradiated
at the U.S.D.A., A.R.S., Subtropical Horticultural Research Station in Miami, Florida.
The eggs were subjected to three doses of gamma radiation from a cobalt-60 source by
the application of a constant amount of radiation for a variable amount of time in min-
utes. Because the cobalt-60 source decayed over the period of the three tests, the 0.075,
0.150, and 0.300 kGy doses were accomplished by exposing the fruit to: 0.16927 kGy/min
in test 1 on 28 September 1988 for 0.443 min, 0.886 min, and 1.772 min, respectively;
0.16742 kGy/in in test 2 on 20 October 1988 for 0.448 min, 0.896 min, and 1.792 min,
respectively; 0.15337 kGy/in in test 3 on 29 June 1989 for 0.489 min, 0.978, and 1.956
min, respectively. These doses are equivalent to those received by carton fruit in com-
mercial radiation facilities (von Windeguth & Ismail 1987). Two controls were used: A
static control (CS), maintained at Lake Alfred, Florida and a mobile control (CM),
transported with the treated eggs.
All eggs were maintained in an environmental growth chamber at 28C, 24 h scotoph-
ase and 100% RH. Mortality was evaluated by counting the number of eggs that did
not produce larvae within 3 weeks after the last of the larvae had emerged from the
controls. The number of eggs that did not produce larvae were divided by the total
number of eggs irradiated to give the mortality percentages. Significant differences
among all treatment and control means were evaluated by general linear model proce-
dures and Duncan's multiple range tests (SAS Institute 1988).


One hundred percent mortality of all eggs occurred with the use of 0.300 kGy (Table
1). There was also 100% mortality of all young (1-5 day) eggs subjected to either 0.075
or 0.150 kGy. Sixteen and 0.1% of older (6-12 day) eggs hatched after being irradiated
with 0.075 and 0.150 kGy, but these amounts were insignificantly different for dose x
age (F = 0.70, df = 4, Pr > F 0.60). Other research has revealed variable susceptibility
among the different developmental stages of an insect species. Cornwell (1966) found
this was true for Ceratitis capitata (Weid). Additionally, different ages of the same
stage were found to be variably sensitive. One to 3-day-old eggs of Gibbium psyllsides
(Czenpinski) were prevented from hatching by cobalt-60 irradiation with only 0.0175
kGy. However, 0.10 kGy were required for 100% mortality of 5-day-old eggs, and 0.40
kGy was necessary for 7-day-old eggs of the same species (Brower 1972).
There was a 75% hatch of controls in trial 3, but only a mean of 35% survival for
controls in trial 1 and 2. However, general linear model procedures established signifi-


Florida Entomologist 73(2)


Dose Egg age No. No. emerging Mortality
(kGy) (days) eggs larvae (%)

0 (CS)2 1-5 539 329 39.0 a3
6-12 381 190 50.1 a
0 (CM)2 1-5 512 241 52.9 a
6-12 333 117 64.9 a
0.075 1-5 629 0 100.0 b
6-12 552 90 83.7 b
0.150 1-5 564 0 100.0 b
6-12 509 1 99.9 b
0.300 1-5 611 0 100.0 b
6-12 519 0 100.0 b

'Data are comprised of total amounts for three tests consisting of at least 50 eggs/replication and three replications/
dose/age/trial except for controls which were replicated once in trial 1, and three times in tests 2 and 3.
2Average percent hatch of all controls = 66.6% (CS) and 42.4% (CM).
'Employing Duncan,s multiple range test (SAS Institute 1988), controls were significantly different statistically
from dose for all three tests at 0.05 level.

cant differences between treatments and controls for all three tests (F1 = 10.01, df =
4, Pr > F = 0.0033; F2 = 7.44, df = 4, Pr > F = 0.001; and F3 = 55.9, df = 4, Pr
> F 0.001 for tests 1, 2, and 3, respectively). The difference between dose and control
for all three tests combined revealed a very significant difference (F = 28.46, df = 4,
Pr > F = 0.0001). Mortality for gamma irradiated eggs showed no significant differ-
ences between tests (F = 0.29, df = 2, Pr > F = 0.75).
Research has not focused only on the effective dosages required for insect eradica-
tion, but also examined the phytotoxic effects of irradiation on foods. Certain trends
have emerged: riper fruits are less prone to injury whereas fruits containing the highest
percentages of water are more susceptible. However, doses of 0.04-0.60 kGy, which are
proven effective in causing 100% mortality of C. capitata in oranges (Citrus sinensis)
(Fesus et al. 1981) and 0.10-0.30 kGy, which controls Anastrepha suspense (Loew) in
grapefruit (Citrus paradisi Macf.) (Burditt et al. 1981), do not significantly alter fruit
quality. Hatton et al. (1982, 1984), quantified rind injury to grapefurit at 1, 4, and 9%,
at doses of 0.075, 0.15, and 0.30 kGy, thus, allowing them to still be marketed. These
results were also reported by von Windeguth (1982). With only a 0.300 kGy dose of
gamma radiation necessary to affect 100% mortality in all ages of Fuller rose beetle
eggs, phytotoxic effects on fruit should not be of concern with this dose also adequate
for the simultaneous eradication of the Caribbean fruit fly in grapefruit (von Windeguth
& Ismail 1987). These results indicate that gamma radiation is one solution for a safe
quarantine treatment for Fuller rose beetle eggs on export citrus.


The authors wish to thank D. von Windeguth for irradiating all three groups of FRB
eggs, Barry Rogers for assistance in the transport and mortality counts of these eggs,
and H. N. Nigg, C. C. Childers, and H. W. Browning for reviewing this manuscript.
We also thank C. Evans, J. Morris, P. Hicks, B. Thompson, W. Tomlinson, and T.
Hardy of the Word Processing Department for the typing and editing of this manuscript.


June, 1990

Coats & Ismail: Ovicidal Effects of Gamma Radiation

This research was supported by grants from the Florida Department of Citrus
(#88044), the California Department of Food and Agriculture (#88-0622, and the
California Citrus Research Board (CRB-256-73-550).
Florida Agricultural Experiment Stations Journal Series No. R-00161.


ANONYMOUS. 1966. Pantomorus cervinus (Boh.). Commonwealth Inst. Entomol.,
Distribution Maps of Pests, Serv. A Map 214: 1-2.
ANONYMOUS. 1988. A race with the calendar. Citrograph 73: 132-135.
BALOCK, J. W., AND L. D. CHRISTENSON. 1956. Effect of gamma rays from Cobalt
60 on immature stages of the "Oriental fruit fly" (Dacus dorsalis Hendel) and
possible application in commodity treatments. Proc. Hawaii Acad. Sci., 31st Ann.
Mtg. p. 18.
BROWER, J. H. 1972. Interaction of age and radiation dosage on hatch of Gibbium
psylloides eggs (Coleoptera:Ptinidae). Ann. Entomol. Soc. Am. 65(5): 1237-1238.
BRYNJOLFSSON, ARI. 1989. Future radiation sources and identification of irradiated
foods. Food Technol. 43(7): 84-89.
BURDITT, A. K., JR. 1982. Food irradiation as a quarantine treatment of fruits. Food
Technol. 36(11): 51-62.
VON WINDEGUTH, AND P. C. SHAW. 1981. Low-dose irradiation as a treat-
ment for grapefruit and mangoes infested with Caribbean fruit fly larvae. U.S.
Dept. Agric. ARS-S-10.
COATS, S. A., AND C. W. McCoY. 1989. Fuller rose beetle, Pantomorus cervinus
(Coleoptera:Curculionidae), ovipositional preference on Florida Citrus. J. Econ.
Entomol. (In press).
CORNWELL, P. B. 1966. Irradiation as a quarantine control measure. Food irradiation.
Proc. Symp. IAEA, Karlsrube. p. 381.
COUEY, H. M. 1983. Development of quarantine systems for host fruits of the medfly.
HortScience 18: 45-47.
FESUS, I., L. KADAS, AND B. KALMAN. 1981. Protection of oranges by gamma
radiation against Ceratitis capitata Wied. Acta Alimentaria 10(4): 293-299.
GJERDE, P. PHILLIPS, AND N. SAKCVICH. 1987. The Fuller rose weevil: Prog-
ress report on a potential disaster. Citrograph 72(8): 147-150.
W. F. REEDER. 1982. Phytotoxicity of gama irradiation on Florida grapefruit.
Proc. Florida State Hort. Soc. 95: 232-234.
VON WINDEGUTH, AND V. CHEW. 1984. Phytotoxic responses of Florida grape-
fruit to low-dose irradiation. J. Am. Soc. Hort. Sci. 109(5): 607-610.
KADER, ADEL A. 1986. Potential applications of ionizing radiation in postharvest
handling of fresh fruits and vegetables. Food Technol. 40(6): 117-121.
M. URBAIN. 1984. Irradiation from CAST 1984-1. Council for Agricultural Sci-
ence andl TechnologY, Ames, Iowa.
KOIDSUMI, K. 1930. Quarantine studies on the lethal action of X-rays upon certain
insects. J. Soc. Trop. Agric., Taiwan. 7: 243.
MATSUYAMA, A., AND K. UMEDA. 1983. Sprout inhibition in tubers and bulbs. pp.
159-213. in E. S. Josephson and M. S. Peterson (eds.), Preservation of Food by
Ionizing Radiation, Vol. III. CRC Press, Boca Raton, Florida.
MCMULLIN, W. H., AND J. G. YEAR. 1982. Working report DOE/USDA/AIBS
workshop on low-dose radiation treatment of agricultural commodities.
Washington, D.C.
SAS INSTITUTE. 1988. SAS user's guide: Statistics. SAS Institute, Cary, North

242 Florida Entomologist 73(2) June, 1990

VON WINDEGUTH, D. L. 1982. Effects of gamma irradiation on the mortality of the
Caribbean fruit fly in grapefruit. Proc. Florida State Hort. Soc. 95: 235-237.
VON WINDEGUTH, D. L., AND M. A. ISMAIL. 1987. Gamma irradiation as a quaran-
tine treatment for Florida grapefruit infested with Caribbean fruit fly, Anas-
trepha suspense (Loew). Proc. Florida State Hort. Soc. 100: 5-7.


Subtropical Horticulture Research Station
Agricultural Research Service
U.S. Department of Agriculture
13601 Old Cutler Road
Miami, Florida 33158


'Marsh' white grapefruit, Citrus paradisi (Macf.). infested with eggs and larvae of
Caribbean fruit fly, Anastrepha suspense (Loew) were subjected to ionizing radiation
at several low doses followed by cold (1.1C) storage for 0 to 8 days. Data analyses
indicated that an irradiation dose of 50 Gray followed by 5 days of cold storage will give
in excess of probit 9 level of quarantine security. A test involving more than 100,000
insects infesting grapefruit confirmed the efficacy of this treatment.


Frutas del cultivar 'Marsh' de toronja blanca infestadas por huevos y larvas de la
mosca frutal del caribe, Anastrepha suspense (Loew) fueron sometidas a irradiaci6n de
ionizaci6n a varias dosificaciones bajas, seguidas por almacenaje frio (1.10C) por periods
de 0 a 8 dias. Analisis de los datos indicaron que una dosificaci6n de irradiaci6n de 50
gray seguida por 5 dias de almacenaje frio producira un nivel en exceso de probit 9 de
seguridad cuarentenal. Un ensayo que envolvi6 mAs de 100,000 insects que infestaron
las frutas de toronja confirm esta observaci6n.

Following the ban by the Environmental Protection Agency (Ruckelshaus 1984) of
ethylene dibromide (EDB) as a fruit fly quarantine treatment for grapefruit, Citrus
paradisi (Macf.), many different treatment methods have been investigated. Benschoter
(1983, 1984, 1987, 1988) reported on the effect of low temperature alone or in combina-
tion with modified atmospheres or fumigants on immature stages of the Caribbean fruit
fly (CFF), Anastrepha suspense (Loew) infesting grapefruit. Benschoter & Witherell
(1984) presented data on the lethal effects of suboptimal temperatures on CFF. Gould
(1988) published on combining hot water and cold storage as a CFF quarantine treatment.
Because some of the proceeding authors had reported a synergistic or enhancing effect
of a given treatment when followed by low temperature (about 1C), we decided to
determine if low dose radiation (<150 Gy) followed by low temperature storage for a

von Windeguth & Gould: Gamma Irradiation

short time (<15 days) would provide an acceptable level of control of CFF without
causing any deleterious phytotoxic effects on the grapefruit. Hatton et al. (1982) re-
ported on the phytotoxic effects of higher doses (75 to 900 Gy) of gamma radiation on
grapefruit. The Gray (Gy) has replaced the formerly used term rad (1 Gy is the equiva-
lent of 100 rad); see footnote 1, table 2 for definition of Gray.
Benschoter (1983, 1984) reported that 1.7C (350F) for 14 days destroyed all the eggs
and larvae of CFF infesting grapefruit, and in a later report (1984) that this temperature
for 15.5 days gave probit 9 security i.e., no more than 32 survivors in 106 treated insects.
The use of ionizing gamma irradiation as a commodity treatment for grapefruit infested
with CFF has been reported by Burditt et al. (1981), von Windeguth (1982), and von
Windeguth & Ismail (1987). An applied dose of 150 Gy was recommended to provide a
probit 9 (99.9968% mortality [Baker 1939]) level of quarantine security. A confirmatory
test of the 150 Gy dose with more than 100,000 immature CFF infesting grapefruit
indicated that based on the development of adult flies from the irradiated immatures,
this dose provided in excess of 99.9968% control of insects.
Although combination treatments require that quarantine officials monitor and con-
trol more parameters, it was felt that there were definite advantages to this particular
combined treatment because of its greater speed and energy saving. Because the radia-
tion output of most commercial irradiators is constant (depending on the number of
placks of source raised), a reduced dose per unit of commodity will shorten the time
that the grapefruit are irradiated. A 50 Gy treatment of commodity would allow almost
3 times as much of the commodity to be treated in the same amount of time as a 150
Gy treatment resulting in a saving of both time and energy.


Commercially harvested and packed grapefruit were obtained from the citrus area
of central Florida. During packing they were washed, waxed, and treated with fun-
gicides. Upon arrival in Miami, the cartons of grapefruit were placd in refrigeration at
10 or 15.5C dependingg on the season) until needed.
Fruits were infested for 1 or 2 weeks depending on the time of the year (longer in
mid-winter, shorter in the warmer season) in an outdoor infestation cage containing
large numbers (approximately 100,000) of gravid ovipositing female CFF. Following
removal from the infestation cage, the fruits were divided into 2 groups. One was
irradiated immediately to assure that some of the immature stages were unhatched eggs,
and the other was held in standard 4/5 bu shipping cartons for one week to assure that
the treatment contained mature 3rd instar larvae and then irradiated.
Irradiations were performed in a Model 220 irradiator (Atomic Energy of Canada
Limited [AECL]) loaded with 14 pencils containing 26,060 curies of Cobalt60 (Jan 1983).
The measured dose rate at the center of the chamber during the time of this study was
from 184.7 to 155.1 Gy per minute, depending on the date when the irradiation was
performed. Because of the small size of the chamber (20.27 x 15.49 cm), only two fruits
were irradiated at a time. Following irradiation, the fruits were packed in cartons with
mold inhibiting biphenyl pads between layers.
The cartons of irradiated fruits were placed on wire shelves in a 1.1C walk-in
refrigerator. Type T thermocouple probes attached to a Fluke model 2285B Data Logger
were placed in several locations inside the fruits, cartons, or room. Temperatures were
recorded at 4-hr intervals. The innermost fruits normally required 48 hrs to cool from
ambient irradiation temperatures (usually around 25C) to the desired cold treatment
temperature. Actual cold treatment time commenced when the fruits had been in the
walk-in refrigerator for 48 hrs. Fruits were removed from the walk-in refrigerator at
2, 4, 6, or 8 days following the start of the cold treatment depending on the test in

Florida Entomologist 73(2)

progress. Following removal from the carton, the fruits were placed in fruit-holding
towers (screened racks which hold approximately 4 cartons of grapefruit, about 144
fruits). Mature larvae that survived the treatments exited the fruit, fell through the
funnel located in the base of the tower and into a sand filled box where they pupated.
The sand was sifted at weekly intervals for up to 6 weeks and the recovered pupae were
held in moist vermiculite at 25C for adult emergence. Control fruits did not undergo
irradiation and cold exposure, but otherwise were treated exactly the same. For each
lot of fruits, about 1/3 were used as control and 2/3 were treated. The number of insects
treated was determined from the mean number of pupae per fruit found in the control
multiplied by the number of fruits subjected to the treatment.
Evaluation of the efficacy of commodity treatments involving ionizing radiation dif-
fers from those using heat, cold, or fumigation in that the latter treatments generally
kill the stage of the insect treated in situ. With radiation, some mortality will occur at
the stage treated, but many larvae will crawl out of the fruits and successfully pupate.
The actual efficacy of the treatment should be based on adult emergence.
A confirmatory test consisting of 13 separate tests with large numbers of infested
fruits in each test, was done over a period of several months. In this confirmatory test,
all fruits were irradiated at the same rate and held at the same temperature and length
of time in the walk-in refrigerator.
To determine if there were any potential phytotoxic effects of the combined treat-
ments, fresh, newly received, unrefrigerated fruits were irradiated at levels within and
exceeding the test rates and refrigerated for times within and exceeding the test times.
Fruits were evaluated for an increase in normal rot and decay as well as observable
defects attributable to the cold or irradiation treatment. For these tests, 4 cartons of
grapefruit (36 fruits/carton) were irradiated at each of the following doses: 50, 100 or
300 Gy. They were held at 1.10C for 3, 6, or 9 days and then compared to unirradiated,
uncooled controls, as well as with each other.


Data on mortality (reduction in pupae per fruit produced) are presented in Table 1.
These mortalities agree closely with those reported by Benschoter (1984) from which
he projected a 15.5 day cold exposure for probit 9 mortality of CFF immatures in
grapefruit. The previously mentioned confirmatory test by von Windeguth indicated
that 150 Gy radiation treatment by itself was sufficient to provide probit 9 quarantine
security. In this test, 13,704 grapefruit containing 114,606 immature insects were ir-
radiated, and no adults emerged from the puparia that were recovered.
For the present series of tests, days of exposure to cold and dos of irradiation were
selected that would, as a minimum, decrease by at least half the cold time and irradiation


Days at No. fruit No. pupae Mean pupae % reduction
1.1C treated recovered per fruit due to cold

0 651 7567 11.624 -
2 324 1472 4.543 60.92
4 324 261 0.806 93.07
6 324 51 0.157 98.65
8 324 5 0.015 99.87


June, 1990

von Windeguth & Gould: Gamma Irradiation

dose. Data in Table 2 are a composite of results of tests performed during 2 citrus
seasons. Natural mortality of the control and treated insects were assumed to be equal
and therefore did not enter into the calculations. The regression procedure of SAS
(1985) was used to obtain a multiple regression of percent mortality on a log/log-scale
of radiation dosage (Grays) and cold storage (days). Percent mortality was transformed
using the arcsin transformation as described by Steel & Torrie (1980). Both radiation
and cold storage effects were significant (Table 3); the r2 value was 0.6273. The regres-
sion equation is given below:

Arcsin sqrt (% mortality) = 0.6902657 + 0.328759 (loglO Grays) +
0.611065 (loglO days cold storage)

If percent mortality is fixed at 99.9968 (probit 9), and the days of cold storage are
held constant, the regression equation can be used to estimate the radiation dose re-
quired to achieve the desired mortality. The same holds true if the dose is held constant
and the days of cold treatment required for probit 9 mortality (Fig. 1) are estimated.
Using the regression equation and data from Table 2, it was estimated that a dose
of 50 Gy and a cold (1.1C) exposure time of 5 days (after a 2-day cooldown period)
would provide in excess of probit 9 quarantine security. Using this projection, a total


Irradiation No. Insects killed % Mortality
Days at dose insects
1.10C (Grays)' treated2 Immatures Adults Immatures Adults

0 5 3808 116 1626 3.0460 42.6996
10 3808 136 2872 3.5710 75.4202
20 3808 2935 3758 77.0745 98.6870
40 3808 3363 3808 88.3141 100
80 3808 3556 3808 93.3824 100
2 5 3097 1501 2564 48.4663 82.7898
10 3097 2238 2983 72.2635 96.3190
20 3097 2674 3096 86.3416 99.9677
40 3097 2563 3097 82.7575 100
80 3097 2723 3097 87.9238 100
4 5 1153 1039 1133 90.1127 98.2654
10 1153 887 1122 76.9297 97.3114
20 1146 1123 1145 98.7784 99.9127
40 1153 1123 1153 97.3981 100
80 1153 1150 1153 97.7398 100
6 5 2828 2804 2825 99.1513 99.8939
10 2828 2816 2826 99.5757 99.9293
20 2845 2839 2844 99.7891 99.9649
40 2845 2842 2845 99.8946 100
80 2863 2863 2863 100 100

'The Gray (Gy) is a unit of energy absorbed from ionizing radiation by matter through which the radiation passes.
A radiation dose of 1 Gy involves the absorption of 1 J of energy by each kilogram of matter through which the radiation
Insects treated is determined by the mean number of pupae per grapefruit from the control multiplied by the
number of fruits treated in its corresponding irradiation/cold treatment. The ratio of control fruits to treated fruits
was about 1:2.


Florida Entomologist 73(2)


Estimate S. E. Prob. > T

Intercept 0.692657 0.123 0.0001
Grays 0.328759 0.077 0.0003
Days of cold 0.611065 0.148 0.0005

of 104,326 immature insects (based on pupae recovered in the controls) infesting 8,592
grapefruits were treated at the 50 Gy/5-day combined treatment rate. A total of 239
pupae were recovered (99.7709% mortality) and 0 adults emerged from these pupae
(100% adult mortality).
Other combinations of gamma irradiation and days of cold could be used, but the 50
Gy/5-day combination gave a 66% reduction in the recommended dose of 150 Gy and
about a 66% reduction in the 15.5 days at 1.7C recommended by Benschoter (1983).
Limited phytotoxicity tests on early (October) and late (March) grapefruit indicated no
obvious visible damage to the fruits at the 50 Gy/5-day treatment rate.
This combination of treatments should provide an acceptable method of quarantine
treatment that could be used for interstate shipment of grapefruit when the commodity
requires treatment because of insect pest restrictions. The treatment required a briefer
period of time for refrigeration and a lower dose of irradiation than either a cold treat-
ment or irradiation alone.




0 1 2 3 4 5 6 7 8 9 10

Fig. 1. Relationship of the dosage of radiation (Gray) and the number of days cold
storage at 1.1F to 99.9968% (Probit 9) mortality.


June, 1990

von Windeguth & Gould: Gamma Irradiation


Mention of a commercial or proprietary product does not constitute an endorsement
by the U.S. Department of Agriculture.


BAKER, A. C. 1939. The basis for treatment of products where fruit flies are involved
as a condition for entry into the United States, USDA Circ. No. 551. 7 p.
BENSCHOTER, C. A. 1983. Lethal effects of cold storage temperatures on Caribbean
fruit fly in grapefruit. Proc. Florida State Hort. Soc. 96: 318-319.
BENSCHOTER, C. A. 1984. Low-temperature storage as a quarantine treatment for
the Caribbean fruit fly (Diptera: Tephritidae) in Florida citrus. J. Econ. Entomol.
77: 1233-1235.
BENSCHOTER, C. A. 1987. Effects of modified atmospheres and refrigeration temper-
atures on survival of eggs and larvae of the Caribbean fruit fly (Diptera: Teph-
ritidae). J. Econ. Entomol. 80: 1223-1225.
BENSCHOTER, C. A. 1988. Methyl bromide fumigation and cold storage as treatments
for California stone fruits and pears infested with the Caribbean fruit fly (Dipt-
era: Tephritidae). J. Econ. Entomol. 81: 1665-1667.
BENSCHOTER, C. A., AND P. C. WITHERELL. 1984. Lethal effects of suboptimal
temperatures on immature stages of Anastrepha suspense. Florida Entomol.
67: 189-193.
WINDEGUTH, AND P. E. SHAW. 1981. Low dose irradiation as a treatment for
grapefruit and mangos infested with Caribbean fruit fly larvae. USDA, ARS,
ARR-S-10. 9 p.
GOULD, W. P. 1988. A hot water/cold storage quarantine treatment for grapefruit
infested with the Caribbean fruit fly. Proc. Florida State Hort. Soc. 110: 190-192.
W. F. REEDER. 1982. Phytotoxicity of gamma irradiation on Florida grape-
fruit. Proc. Florida State Hort. Soc. 95: 232-234.
RUCKELSHAUS, W. D. 1984. Ethylene dibromide, amendment of notice of intent to
cancel registration of pesticide products containing ethylene dibromide. Federal
regist. 49(70): 14182-14185.
SAS INSTITUTE. 1985. SAS Users Guide: Statistics. SAS Institute, Cary, NC. p.
STEEL, R. G. D., AND J. H. TORRIE. 1980. Principles and Procedures of statistics.
McGraw-Hill, New York.
VON WINDEGUTH, D. L. 1982. Effects of gamma irradiation on the mortality of the
Caribbean fruit fly in grapefruit. Proc. Florida State Hort. Soc. 95: 235-237.
VON WINDEGUTH, D. L., AND M. A. ISMAIL. 1987. Gamma irradiation as a quaran-
tine treatment for Florida grapefruit infested with Caribbean fruit fly, Anas-
trepha suspense, (Loew). Proc. Florida State Hort. Soc. 100: 5-7.


Florida Entomologist 73(2)


Department of Zoology
Michigan State University
East Lansing, Michigan 48824


A new species, Ptenothrix (Ptenothrix) austalis is described from Georgia, South
Carolina and Florida. The species can be identified by color pattern and chaetotaxy.
Dicyrtoma (Dicyrtomina) rossi (Wray) is redescribed from new material. Evidence is
presented that may lead to placing this species in a new subgenus. New collection
records for 3 little known species are presented: Ptenothrix (Dicyrtoma) fusca (Lub-
bock), Ptenothrix (Ptenothrix) macomba Way and Ptenothrix (Ptenothrix) curvilineata


Este papel es parte de una series que describe Col6mbolas de la plant del rio Savan-
nah, en Aiken, Carolina del Sur y de los estados alrededor del sudeste. El prop6sito es
el describir una especie nueva, re-describir otra y presentar la extension de la distribu-
ci6n de 3 species raras.

This paper is part of a series describing Collembola from the Savannah River Plant,
Aiken, South Carolina and surrounding southeastern states. The purpose is to describe
a new species, redescribe another and present range extensions of 3 rare species.

Ptenothrix (Ptenothrix) australis NEW SPECIES

COLOR: Background cream to white with purple and olive pigment laid down in
irregular polygons. Head with interocular maculae dark purple; frons below antenna
bases with 2 + 2 dark purple maculae, grooves darker becoming lighter at edges; gena
and post genal area with purple polygons; ANT I distally purple, dark basally; ANT II
light, purple confined to central area of segment; ANT III with purple on basal half and
on apical portion; ANT IV with basal half white, distally purple. Body with first thoracic
segment outlined in dark purple, other thoracic segments with irregular maculae; great
abdomen with 2 lateral dark purple patches, o.e dorsal, the other ventral followed by
a field of olive purple interspersed with white rings; lesser abdomen light purple, lower
valves darker; legs with purple bands beyond trochanter; furcula with light purple
dusting (Figs. 1 & 2).
HEAD: Eyes 8+8; ocelli A & B subequal, C, D & E 1/2 diameter of A; F, G & H
slightly smaller than A, subequal (Fig. 3). Mean antennal ratio 2:5:5:2; ANT III divided
into 8 distal subsegments, subapical sensilla exposed, lying in shallow depressions; ANT
IV superficially divided into 8 subannulations. Outer maxillary lobe consists of simple
palp and 1 sublobal hair (Fig. 4). labrum with setal pattern 6/5,2,4 (Fig. 5). Dorsal setae
of head (B E) short, spine-like; 5 spine-like, unpaired facial setae; 1+1 oval organs
on lower gena (Fig. 6). FORELEG: Coxa with 1 seta and no oval organ (Fig. 7);


June, 1990

Snider: Southeastern U.S. Dicyrtomidae

1 2

Ptenothrix australis n. sp. Fig. 1 & 2 Habitus.

trochanter with 2 anterior and 2 posterior setae, distal anterior seta reduced to a setula
(Fig. 8); femur with basal posterior and distal anterior oval organs, cup sensillum on
outer margin (Fig. 9); tibiotarsus with 4 cup sensilla and 3 oval organs (almost courte
epines") on anterior surface (Fig. 10), lacking an oval organ on posterior surface (Fig.
11); pretarsus with anterior and posterior setulae; unguis lack tunica, with basal lateral
teeth, 2 inner teeth well developed; unguiculus with corner tooth, subapical filament
reaching beyond tip of unguis (Fig. 12). MESOLEG: coxa with 3 anterior setae and 1
courte epine" (Fig. 13); trochanter with 4 anterior and 1 posterior setae (Fig. 14);
femur with basal posterior and distal anterior courte epines", cup sensillum on outer
margin (Fig. 15); tibiotarsus with 5 cup sensilla and 3 courte epines" on anterior surface
(Fig. 16), 1 courte epine" on posterior surface (Fig. 17); pretarsus with anterior and
posterior setulae; unguis similar to foreclaw, unguiculus with corner tooth, subapical
filament reaching beyond tip of unguis (Fig. 18). METALEG: Coxa with 4 anterior setae
and 1 courte epine" (Fig. 19); trochanter with 5 anterior and 1 posterior setae (Fig.
20); femur with basal posterior oval organ, distal anterior courte epine", and outer cup
sensilla (Fig. 21); tibiotarsus with 5 cup sensillum and 3 courte epines" on anterior
surface (Fig. 22), posterior surface with 2 strongly knobbed, serrate differentiated
setae (Fig. 24) and no courte pines" (Fig. 23); pretarsus with anterior and posterior
setulae; unguis similar to other claws; unguiculus with corner tooth, and subapical fila-
ment reaching beyond tip of unguis (Fig. 25). GREAT ABDOMEN: Collophore with 1+1
subapical and 1+1 lateral setae (Fig. 26). Corpus of retinaculum with 6 setulae, ramus
with 3 teeth and basal horn (Fig. 27). Manubrium with 9 + 9 dorsal setae (Fig. 28). Dens
with 3-2-1-1......1 Ve setae (Fig. 29), dorsal E and L setae finely serrate (Fig. 30), E1/E2
= 1.6 and E3/E2 = 2.2 (Fig. 31). Mucro with inner and outer teeth (Fig. 32).
LESSER ABDOMEN: Circumanal setae M,M' and N spine-like, smooth, seta sa seta-
like; setal pattern, expressed as: M M' N T H G Ao sa (Fig. 33);
subanal appendage distally curving, acuminate (Fig. 34). BODY CLOTHING: Anterior
body setae (AA-EE) seta-like, smooth, posterior dorsal setae short, dagger-like.
Parafurcular setae 11:9, undifferentiated, 2 cup sensilla (Fig. 35). Length up to 2mm.
DIAGNOSIS: Ptenothrix (Ptenothrix) australis n.sp. keys out in Christiansen and
Bellinger (1981) to D. (Dicyrtoma) hageni (Folsom). These authors attempted to synthe-
size a taxonomic key to species without first separating the family Dicyrtomidae into
tribes as proposed by Richards (1968). The result is an unclear separation of Dicyrtoma
and Ptenothrix as new species are uncovered. The two genera can easily separated by
presence of bothriothrix D and 2 differentiated setae on the metatibiotarsus of Pteno-
thrix spp. and lack of D and presence of 3 spines on posterior surface of the metatibiotar-
sus of Dicyrtoma spp. Further, Dicyrtoma hageni has a distinctive color pattern that

Florida Entomologist 73(2)









22 23


< ^
S\ I
N *



k F
Ah /\

II / -
/ *\ '

1\/ 0 11
10 11

Fig. 3. eyepatch. Fig. 4. outer maxillary lobe. Fig. 5. labrum. Fig. 6. setal pattern
of head. Fig. 7. forecoxa. Fig. 8. foretrochanter. Fig. 9. forefemur. Fig. 10. foretibiotar-
sus, anterior surface. Fig. 11. foretibiotarsus, posterior surface. Fig. 12. foreclaw. Fig.
13. mesocoxa. Fig. 14. mesotrochanter. Fig. 15. mesofemur. Fig. 16. mesotibiotarsus,
anterior surface. Fig. 17. mesotibiotarsus, posterior surface. Fig. 18 mesoclaw. Fig. 19.
metacoxa. Fig. 20. metatrochanter. Fig. 21. metafemur. Fig. 22. metatibiotarsus, an-
terior surface. Fig. 23. metatibiotarsus, posterior surface. Fig. 24. differentiated setae
of metatibiotarsus. Fig. 25. metaclaw.




June, 1990


Snider: Southeastern U.S. Dicyrtomidae 251

u 27 V LA- k I

32 ,/ !< /-',

31 32 30 29
34 .

(/ 33 35

38 40 41
38 41
43 \



46 47 48
Fig. 26. collophore. Fig. 27. retinaculum. Fig. 28. manubrium. Fig. 29. dens, ventral
surface. Fig. 30. dens, dorsal surface. Fig. 31. dorsal E setae. Fig. 32. mucro. Fig. 33.
circumanal setae. Fig. 34. subanal appendage. Fig. 35. parafurcular setae. Fig. 38.
eyepatch. Fig. 39. antennal segment I. Fig. 40. outer maxillary lobe. Fig. 41. labrum.
Fig. 42. head, facial setae. Fig. 43. forecoxa and coxal organ. Fig. 44. foretrochanter.
Fig. 45. forefemur. Fig. 46. foretibiotarsus, anterior surface. Fig. 47. foretibiotarsus,
posterior surface. Fig. 48. foreclaw.

Florida Entomologist 73(2)

36 37

Dicyrtoma (Dicyrtomina) rossi (Wray) Figs. 36 & 37. habitus.

will easily separate it from the new species. Ptenothrix (Ptenothrix) marmorata Pack-
ard) has spine-like circumanal setae arranged in a similar pattern to australis. The two
species can be distinguished by pattern and presence of 5 unpaired facial setae on
australis, while marmorata has 2.
TYPES: Holotype (9), on slide and 1 paratype in alcohol; Georgia, Rabun County,
Tallulah Falls, gorge, leaf litter, "March 1, 1981," D. Yehling, collector. South Carolina,
Aiken County Savannah River Plant, rail bridge on Road F, soil from tree stump,
"August 30, 1984," Barnwell County, SRP, along creek off Road B, litter, "November
8, 1983", W. W. Hargrove, collector 1 adult. Florida, Dade County, Everglades National
Park, near "Cottonmouth Pond, litter. "March 22, 1982", W. W. Hargrove, 4 subadults.
All specimens deposited in the Department of Entomology Museum, Michigan State

Dicyrtoma (Dicyrtomina) rossi (WRAY), 1952

COLOR: Background white or cream with purple and blue pigment laid down in
mosaics. HEAD: with broken median interocular stripe, blue maculae below antennal
bases; lower frons with median stripe, distally spreading over labral area, gena laterally
with 3 irregular, diagonal lines; ANT I with dark purple distally; ANT II lightly dusted
with blue-purple; ANT III dark blue, darker distally; ANT IV dark blue. BODY: thoracic
area dorsally with 2 paramedial lines, laterally with broken bands in precoxal area;
abdomen with lateral, irregular band, postero-dorsal area with horseshoe shape band,
dorsal macula dark blue-black (in life abdominal banding chestnut-brown). LESSER
ABDOMEN: With median, basal blue-black macula, distally with blue mosaics. Parafur-
cular area with blue patches. APPENDAGES: Legs with light blue banding, becoming
lighter distally. Furcula with blue macula distally on manubrium, dens with light blue
dusting (Figs. 36 & 37).
HEAD: Eyes 8+8, ocelli C and D 1/2 diameter of others (Fig. 38). Mean antennal
ratio (fresh mounts) 1:5:6:1.5; ANT I with large basal segment (Fig. 39); ANT III & IV
not subannulated. Outer maxillary lobe consists of simple palp and 1 sublobal hair (Fig.
40). Labrum with setal pattern: 6/7(1),2,4 (Fig. 41). Cranial setae short, seta-like; 6
unpaired facial setae; 2 + 2 oval organs on lower genae (Fig. 42). FORELEG: Coxa with
1 seta and coxal organ, no oval organ (Fig. 43); trochanter with 2 anterior and 2 posterior


June, 1990

Snider: Southeastern U.S. Dicyrtomidae



51 I


1' r -' "


9 67

64 65


Fig. 49. mesocoxa. Fig. 50. mesotrochanter. Fig. 51. mesofemur. Fig. 52.
mesotibiotarsus, anterior surface. Fig. 53. mesotibiotarsus, posterior surface. Fig. 54.
mesoclaw. Fig. 55. metacoxa. Fig. 56. metatrochanter. Fig. 57. metafemur. Fig. 58.
metatibiotarsus, anterior surface. Fig. 59. metatibiotarsus, posterior surface. Fig. 60.
metaclaw. Fig. 61. collophore. Fig. 62. retinaculum. Fig. 63. manubrium. Fig. 64. dens,
ventral surface. Fig. 65. dens, dorsal surface. Fig. 66. dorsal E setae. Fig. 67. mucro.
Fig. 68. circumanal setae. Fig. 69. subanal appendage. Fig. 70. posterior dorsal abdom-
inal setae. Fig. 71. parafurcular setae.



58 59



S 66



254 Florida Entomologist 73(2) June, 1990

setae, distal posterior spur present (Fig. 44); femur with basal and distal oval organs,
cup sensillum on outer edge, 5 anterior and 6 posterior setae (Fig. 45); tibiotarsus with
3 cup sensilla and 1 oval organ on anterior surface (Fig. 46), oval organ present on
posterior surface (Fig. 47); pretarsus with anterior and posterior setulae; unguis with
a tunica and pseudonychia, 2-3 strong lateral teeth and 2 inner teeth; unguiculus without
corner tooth, subapical filament reaching beyond tip of unguis (Fig. 48). MESOLEG:
Coxa with 3 setae and 1 courte epine" (Fig. 49); trochanter with 4 anterior and posterior
setae (Fig. 50); femur with basal and distal oval organs, 9 anterior and 5 posterior setae,
outer cup sensillum present (Fig. 51); tibiotarsus with 5 cup sensilla, no courte epine"
on aterior surface (Fig. 52), 1 courte epine" on posterior surface (Fig. 53); pretarsus
with anterior ard posterior setulae; unguis with strong lateral tooth, tunica,
pseudonychia and 2 inner teeth; unguis with outer lamella reaching beyond tip of unguis,
corner tooth present (Fig. 54). METALEG: Coxa with 4 setae and 1 courte epine" (Fig.
55); trochanter with 5 anterior setae (Fig. 56); femur with basal and distal oval organs,
9 anterior, 2 posterior setae, and 2 posterior setulae, 1 outer margin cup sensillum (Fig.
57); tibiotarsus with 5 cup sensilla on anterior surface (Fig. 58), posterior surface with
2 smooth, slightly knobbed, differentiated setae and 2 courte epines" (Fig. 59); pretar-
sus with anterior and posterior setulae; unguis with 2 lateral teeth, tunica, pseudonychia
and 2 inner teeth; unguiculus with corner tooth, outer lamella just reaching tip of unguis
(Fig. 60). GREAT ABDOMEN: Collophore with 1+1 subapical and 1+1 lateral setae
(Fig. 61). Corpus of retinaculum with 4 distal setulae, ramus with 3 teeth and basal
horn (Fig. 62). Manubrium with 9+9 dorsal setae (Fig. 63). Dens with 4,2,1.....1 Ve
setae (Fig. 64), dorsal E and L setae smooth (Fig. 65), El/E2+ 1.0 and E3/E2= 1.5 (Fig.
66). Mucro with inner and outer teeth (Fig. 67). LESSER ABDOMEN: Circumanal setae
M,M';Ao, sa all seta-like (Fig. 68); subanal appendage curving distally, acuminate (Fig.
69). BODY CLOTHING: Anterior body setae not modified, posterior dorsal setae short,
dagger-like (Fig. 70). Parafurcular setae 8:6, undifferentiated, 2 cup sensilla (Fig. 71).
Length up to 1mm.
DIAGNOSIS: Dicyrtoma (Dicyrtomina) rossi (Wray) was described as having a color
pattern on the body with, "irregular purple pigmented streaks; with a horseshoe-shaped
streak on postero-lateral margin. Dorsally with dark streak between eyes extending
down to front. Purple pigment dispersed on cheeks and vertex of head". Christiansen
and Bellinger (1981) only saw a poor type specimen subadultt) and 2 specimens taken
from Massachusetts. Their color description states, "background color yellow; pigment
when present purplish, usually in stripes on body." The specimens taken in Florida
strongly agree with the description presented by Wray (1952).
Christiansen and Bellinger (1981) state that Dicyrtomina can be distinguished from
other dicyrtomids by a well developed tunica and an undivided third antennal segment.
Betsch (1980) based his diagnosis of genera on Richards (1968). He separated the genus
by absence of bothriothrix D, claws with swollen tunica and pseudonychia, 3 differen-
tiated setae on metatibia, presence of bothriothrix A, parafurcular setae undifferen-
tiated, many setae on head and anterior of great abdomen spiniform. In his definition
of Dicyrtomina, the number of differentiated tibiotarsal setae, head and body setae,
and parafurcular setae are of great importance.
Among the 3 North American species, only Dicyrtoma (Dicyrtomina) minute (Fab-
ricius, 1783) fulfills the criteria for Betsch's (1980) generic designation. Dicyrtoma (Dicyr-
tomina) opalina (Folsom, 1896) has 3 differentiated tibiotarsal setae, but lack the spine-
like head and body setae. Dicyrtoma (Dicyrtomina) rossi has only 2 differentiated setae
and also lacks spine-like body setae. Christiansen and Bellinger (1981) said D. (Dicyr-
tomina) opalina clearly falls within the subgenus Calvatomina Yosii (1966). If they are
correct, and it appears obvious, then we can eliminate D. opalina from the subgenus
Dicyrtomina. This leaves D. (Dicyrtomina) rossi for repositioning.

Snider: Southeastern U.S. Dicyrtomidae




Figs. 72 & 73. Dicyrtoma (Dicyrtoma) fusca (Lubbock)

D. (Dicyrtomina) rossi not only lacks spine-like setae on the head and body, but has
only 2 differentiated tibiotarsal setae. In addition, the parafurcular setae are normal.
In the case of Calvatomina sp. some of these setae are short and spiniform. It becomes
clear that D. (Dicyrtomina) rossi represents a transitional genus between Ptenothricini
and Dicyrtomini. A similar situation exists for Dicyrtomina turbotti Salmon (1948) from
New Zealand. Betsch (1980) could only find 2 differentiated tibiotarsal setae and no
bothriothrix D. However, the specimen he examined was subadult and lacked a tunica,
which may be a condition at early instars. Perhaps when generic revision is undertaken,
these two species will be placed in a separate genus.
COLLECTIONS: 5 specimens on slides and 6 specimens in alcohol deposited in the
Department of Entomology Museum, Michigan State University. Florida, Palm Beach
County, Jupiter Island Park (Bert Reynolds Park), dry grass, sweeping, "December
27, 1982", R. J. & R. M. Snider, collectors. Other records: Illinois and Massachusetts.


Dicyrtoma (Dicyrtoma) aurata (Mills), 1934
Virginia, Giles County, Mountain Lake, elev. 3000', grass sweeping, "August 6,
1981", S. J. Loring & R. M. Snider, collectors. Other collections: Iowa, Michigan, and
North Carolina.

Dicyrtoma (Dicyrtoma) fusca (Lubbock), 1873
(Figs. 72 & 73)

Florida, Volusia County, 2.5 miles S of Oak Hill exit on 1-95, grass sweeping, "June
4, 1982," D. Mallow, collector. Other collections: Nova Scotia.

Ptenothrix (Ptenothrix) macomba Wray, 1967
(Figs. 74 & 75)
Georgia, Rabun county, Tallulah Falls, Panther Creek, HWY 17, leaf litter, "March
22, 1981," L. R. Boring, collector; Tallulah Falls, gorge, leaf litter, "March 1, 1981," D.
Yehling, collector; leaf litter, "March 7, 1981", R. J. Snider, collector. North Carolina,
Macon County, Coweeta Hydrology Laboratory, Watershed 13, hardwood litter,


Florida Entomologist 73(2)





Figs. 74 & 75. Ptenothrix (Ptenothrix) macomba Wray.

"November 21, 1980," T. Seastedt, collector; Watershed 10, below weir, cove hardwood
litter, Shope Fork, "January 22, 1981", L. R. Boring, collector. Other collections: Il-

Ptenothrix (Ptenothrix) curvilineata (Wray), 1949
(Fig. 76 & 77)
Virginia, Giles County, Mountain Lake, Mt. Lake Lodge, grass sweeping, "August
6, 1981", R. J. & R. M. Snider, and S. J. Loring, collectors. Other collections: North

Grateful thanks are offered to the Savannah River Ecology Laboratory of the Uni-
versity of Georgia for support provided by contract EY-76-C-09-0819, NERP Program



Figs. 76 & 77. Ptenothrix (Ptenothrix) curvilineata (Wray).

June, 1990


Cassani et al.: Sphenarches anisodactylus in Florida

of the U.S. Department of Energy. This manuscript was reviewed by Drs. Kenneth A.
Christiansen of Grinnell College and Peter F. Bellinger of California State University,
Northridge. Their comments are very much appreciated. The habitus sketches were
rendered by Peter Carrington of Michigan State University.


BETSCH, J. M. 1980. Elements pour une monographie des Collemboles Symphypl
ones (Hexapodes, Apterygotes). Mem. Mus. Nat. Hist., Ser. A, (116): 1-227.
CHRISTIANSEN, K. A., AND P. F. BELLINGER. 1981. The Collembola of North
America North of the Rio Grande, part 4: Families Neelidae and Sminthuridae.
Grinnell College, Grinnell, Iowa: 1043-1322.
FABRICIUS, 0. 1783. Beskrivelse over nogle lidet bekiendte podurer, og en besonder-
lig loppe. K. Danske Vidensk. Selsk. Skr., (2): 296-311.
FOLSOM, J. W. 1896. Two new species of Papirius. Canadian Ent., (28): 119-121.
LUBBOCK, J. 1873. Monograph of the Collembola and Thysanura. London: Ray Soci-
ety, 276 p.
MILLS, H. B. 1934. A Monograph of the Collembola of Iowa. Collegiate Press, Ames,
Iowa, 143 p.
RICHARDS, W. R. 1968 Generic classification, evolution, and biogeography of the
Sminthuridae of the world (Collembola). Mem. Ent. Soc. Canada, (53): 1-54.
SALMON, J. T. 1948. Collembola from the three Kings Islands with a description of
Proisotomina new genus. Rec. Akld. Inst. & Mus., (3)4-5: 291-300.
WRAY, D. L. 1949. Some new Dicyrtoma and key to known species of the United
States (Collembola, Sminthuridae). Bull. Brooklyn Ent. Soc., 44(2): 61-68.
S1952. Some new North American Collembola. Ibid., 47(4): 95-106.
1967. Some new North American Collembola, Ent. News, (78): 227-232.
YosII, R. 1966. On some Collembola of Afghanistan, India and Ceylon, collected by
the Kuphe-Expedition, 1960. Res. Kyoto Univ. Scient. Exped. Karakoran &
Hindukush, 1955, (8): 333-405.


Lee County Hyacinth Control District
P. O. Box 06005
Ft. Myers, Florida 33905

University of Florida, IFAS
Department of Entomology and Nematology
Gainesville, Florida 32611-0711


The life history of Sphenarches anisodactylus (Walker) in Florida is presented.
Descriptions and durations of life stages are given and bionomics relative to the Florida
host, Thalia geniculata (Marantaceae) are discussed. Development time of laboratory










Figs. 74 & 75. Ptenotkrix (Pten othrix) macomba Wray.
"November 21, 1980," T. Seastedt, collector; Watershed 10, below weir. cove hardwood
litter, Shope Fork, "January 22, 1981", L R. Boring, collector. Other collections: Il-
Ptenoifrix (Ptenothrix) eurvilineata (Wray), 1949
(Fig. 76 & 77)
Virginia, Giles County, Mountain Lake, Mt. Lake Lodge, grass sweeping, "August
6, 1981". J, & RK M. Snider, and S. J. Loring, collectors. Other collections: North
Grateful thanks are offered to the Savannah River Ecology Laboratory of the Uni-
versity of Georgia for support provided by contract EY-76-C-09-0819, NERP Program




Figs. 76 & 77. Ptenothrix (PteIrwohrix) cunjifi.neala (Wray).

Florida Entomologist 73(2)

reared individuals from oviposition to adult emergence was 21.8 days for males and 22.5
days for females. There are four larval instars. Adults lived 14-19 days.


Se present el ciclo biol6gico de Sphenarches anisodactylus (Walker) en Florida. Se
proprcionan descripciones y duraciones de los diferentes estadios; se discute la
bion6mica relative al hospedero de la Florida, Thalia geniculata (Marantaceae). El
tiempo de desarrollo de los individu6s bajo condici6nes de laboratorio, desde oviposici6n
hasta el estado adulto, fue de 21.8 dias para machos y 22.5 dias para hembras. Las
larvas pasan por 4 estadios. Los adults vivieron de 14 a 19 dias.

The Pterophoridae, or plume moths, are a mostly inconspicuous group of approx-
imately 600 species worldwide (Stanek & Turner 1977). Twenty genera and 146 species
have been identified from North America (Munroe 1983) with 29 species occurring in
Florida (Kimball 1965). Plume moths are leaf skeletonizers, flower feeders or stembor-
ers. Neunzig (1987) summarized information on pterophorid larvae and illustrated seven
species. Immature stages of most North American plume moth species are unknown.
There are several species of economic importance in the United States some occasionally
causing serious damage. The most important of these is the artichoke plume moth,
Platyptilia carduidactyla Riley, which is a pest of cultivated globe artichokes in Califor-
nia (Lange 1950). More biological information is available on this species than any other
pterophorid in North America.
During a survey of insects occurring on the host plant Thalia geniculata L., a
pterophorid larva was discovered feeding on the flowers. Reared adults were identified
as Sphenarches anisodactylus (Walker) based on comparison of genitalia with the illus-
trations in Adamczewski (1951) and Yano (1963) as well as with identified material in
the United States National Museum (USNM). Our larval material from T. geniculata
is also identical to preserved USNM specimens from pigeon pea (Cajanus cajan) and
matches illustrations and descriptions by Yano (1963) and those by Cotes (1891) under
the name Sphenarches caffer (Zeller).
Larvae of S. caffer were reported tunneling in pods of Dolichos lablab in India
(Cotes 1891). Walsingham, who identified Cotes' specimens, gave the distribution as
West Africa (on "calabash"), Australia, Asia, as well as New Hebrides and Tonga Island
in the Pacific (Cotes, 1891). Yano (1963) reports two varieties of calabash as hosts,
Lagenaria leucantha Rosby var. clavata and gourda. Forbes (1930) listed it as generally
occurring in the Old World tropics and reported Caperonia and pigeon pea as hosts in
Puerto Rico and the West Indies. Bruner et al. (1975) listed it as a pest of pigeon pea
in Cuba.
The most recent revisionary study which included this genus is Adamczewski (1951).
Prior to Adamczewski's work, authors including Cotes (1891), Walsingham (1897),
Meyrick (1910), and Forbes (1930), treated S. anisodactylus (Walker) 1864 as a synonym
of S. caffer (Zeller) 1852). Although the name caffer is older, Adamczewski applies the
name anisodactylus to all the records, specimens, and biological information previously
considered under caffer. Without having examined Zeller's holotype of caffer (located
at the Stockholm Museum, Sweden) he applied the name caffer to a single male specimen
from Natal, considering it a topotype of caffer. Although we suspect anisodactylus is
indeed a synonym of caffer, until all the types can be examined we must use the name
anisodactylus according to Adamczewski.
Sphenarches anisodactylus is previously unreported from the United States. The
only other Sphenarches in North America is S. ontario (McD.), which occurs in Ontario,

June, 1990

Cassani et al.: Sphenarches anisodactylus in Florida

Canada (McDunnough 1927). While the life history of S. anisodactylus is known from
leguminous hosts in the tropics, the following gives a detailed account of the life history
on a completely unrelated host. In this case, both host and insect range from the tropics
of extreme south Florida, to the more temperate areas of central and northeast Florida.


Field observations of S. anisodactylus on Thalia were made at least monthly from
May 1979 to August 1980 and periodically thereafter through 1983 at three locations in
Lee County, Florida. Other collections were made in Glades, Collier, Hendry, Broward
and St. Johns counties. Field observations were made to determine seasonal occurrence
and general abundance as related to host plant condition and flowering.
Sphenarches anisodactylus was also studied in the laboratory during 1982 to deter-
mine life stage characteristics and related behavior. During October 1982, 25-30 field
collected pupae were returned to the laboratory and placed in a one cubic foot cage in
a covered outdoor area. Emerging adults were provided with honey water and allowed
to mate for an 8-12 day period. After mating, 5 females and 3 males were brought into
the laboratory and held individually in plastic cylinders (22 cm high, 11 cm wide) with
screened tops to determine longevity and fecundity. Females oviposit primarily on
flower buds. A fresh flower bud was provided daily to each female and the number of
eggs oviposited was recorded. Mean maximum and minimum temperatures in the labo-
ratory during the study were 25.2 + 0.6 and 23.4 0.8. The photoperiod was main-
tained at 12 h for specimens held inside the laboratory.
The remaining adults were held together in the emergence cage in the laboratory.
Fresh flowers for oviposition were provided as needed. After oviposition, 15 eggs, all
oviposited on the same day, were removed with a camel hair brush and each placed on
individual flowers in clear plastic dishes (8.8 cm diameter; 1.8 cm deep) to determine
instar duration. A square (4 cm x 4 cm) piece of paper towel was moistened and placed
in each dish to maintain humidity. After eclosion, a fresh flower and clean paper were
provided daily.
Another group of 100-150 eggs were held together in an aluminum pan (22 cm diam-
eter; 3.8 cm deep) with a glass cover and fresh flowers provided as needed to feed
emerging larvae. The larvae from pan held eggs as well as some field collected larvae
were used for head capsule measurements and overall body length. All measurements
of larvae were based on specimens killed in boiling water and preserved in 70% ethyl
alcohol. Head capsule measurements were made using a dissecting microscope equipped
with a calibrated ocular grid. Most measurements were made at 40X. Adult measure-
ments were made on pinned specimens. In larval and pupal descriptions, thoracic and
abdominal segments are designated by the prefix T and A, respectively, followed by
the segment number i.e. Tl, A4, etc. General setal nomenclature follows Stehr (1987).
Setal nomenclature of the labrum follows Heinrich (1916).


EGG. (Fig. 1). Eggs are greenish white initially, later becoming pale amber. Egg shape
is elongate-oval with a mean length and width of 0.48 mm + 0.02 mm and 0.31 mm +
0.01 mm respectively (n= 20). The chorion surface appears smooth but SEM examina-
tion at 160X reveals regular patterns of hexagonally arranged aeropyles. Chorionic
ridges are absent. At magnification over 1500x, the surface appears rough and scaly.
The micropyle is a pore-like opening at one end, which is slightly ribbed and surrounded
by outwardly radiating ridges.
LARVA. (Figs. 2-5). Early first instar larvae are pale green. Older larvae (instars 2-4)


260 Florida Entomologist 73(2) June, u19

Fig. 1. Sphenarches anisodactylus ova: scanning electron micrograph at 160x. De-
bris of surface outlines zone of attachment to host plant.

usually become the color of the plant tissue they have been feeding on. Larvae that feed
heavily on flower petals are distinctly rose purple while those that feed on the ovary
or bracts are green. Mature larval length: 10-11 mm. Generally, the prepupa becomes
pale green as feeding ceases. The following description is based on mature larvae.
Head. (Fig. 3). Hypognathous. Six stemmata with stemma 6 equidistant to 4 and to
5, stemma 4 and 5 usually closer to each other than to 3 or 6. Seta S1 posterior to
stemma 2 and equidistant between stemma 1, 3, and 6. S2 posterior to stemma 1, seta
base similar distance from stemma 1 as stemma diameter. S3 ventral to stemma 6 and
posterior to stemma 5, slightly closer to stemma 6. Mandibles with five teeth (Fig. 4).
Labral notch V-shaped extending 1/3 of way to base. Adfrontals reach epicranial notch;
frontoclypeus extends 3/4 way to epicranial notch. F1 setae distinctly above Fa
punctures, AF2 setae slightly posterior of P2 setae and about halfway between front
and epicranial notch. AF1 seta slightly anterior to P1 seta. Labrum with seta M2
directly dorsal of M3. Seta M1 slightly ventral to M2. Seta L2 near ventral margin, L1
and L3 more dorsal and along lateral margin.
Thorax. Prothoracic shield sometimes with dark indentation between D1 and D2
setae; narrow dark margin along light mid-dorsal stripe. XD2 about equidistant from
XD1 and SD1. D1 and SD2 setae spatulate. L group trisetose with posterior seta
spatulate. SV group bisetose. Spiracle conical with collar-like sclerite.
T2 and T3 with trisetose L group on same pinaculum; L3 spatulate and less than 1/2
length of L1 and L2, SV bisetose, D1 spatulate, D2 slightly posterior and ventrad, D2
on T3 less than 1/3 length of D2 on T2. SD1 and SD2 on same pinaculum, SD2 spatulate
and 1/3 length of SD1 on T3, 2/3 length SD2 on T2. Four to five short capitate setae on
edge of or near D pinaculum. One short capitate seta behind D pinaculum 1/2 way

_ __I_~


Cassani et al.: Sphenarches anisodactylus in Florida


3 4 5

Fig. 2-5. Sphenarches anisodactylus mature larva: 2. larva; 3. head; 4. mandible; 5.
dorsal pinaculum on A6.

between pinaculum and posterior margin of segment. T2 with about 8 very short capi-
tate setae along posterior margin of segment.
Abdomen. Scattered very short capitate secondary setae on abdominal segments.
D1 and D2 pinacula fused (Fig. 5) but with a furrow between setae, D1 spatulate
extending anteriorly, D2 twice as long as Dl, 1-4 short capitate setae on edge of common
pinaculum. SD setae spatulate on a common pinaculum; A1-A7 with SD2 1/3 to 1/2
length of SD1 and less broadly spatulate; A8 with SD2 minute and capitate. One small
capitate seta directly posterior to SD pinacula on A1-A7 but not on A8. SD pinacula on
A2-6 with a short capitate seta between SD setae. L1 and L2 pinacula joined, slightly
further from spiracle than SD pinaculum on Al, equidistant on A2; closer to spiracle
than SD pinacula on A3-8. Al-8 with L3 remote, 1 1/2 to 2x as far from L1 and L2 than
from SV pinaculum. L1 and L2 horizontally arranged on A8, a small capitate seta
posterad on a pinaculum. SV group bisetose on Al, A7, trisetose on A2-6, unisetose on
A8. V1 small, midventral distance between about 1/2 distance to SV on Al. 2 V1 near
base of proleg on A3-6. Anal shield with 8 pair of setae, the most anterior and lateral
pair spatulate. Anal proleg with at least 9 setae, slightly less peg-like than abdominal
prolegs. Prolegs on A3-6 long and peg-like about 4X longer than wide. Seven to nine
crochets in a uniordinal mesopenellipse, broadly open laterally.
PUPA. (Figs. 6-7). Length about 9 mm. Pale green initially, later becoming brown.
Body surface except posterior 1/3 to 1/2 of A3/7 finely shagreened with thin transverse
striations. Labial palps small. Maxilla 3/4 length of T1 legs. Antenna nearly as long as

Florida Entomologist 73(2)

Figure 6-7. Sphenarches anisodactylus pupa: 6. dorsal view; 7. lateral view.

T1 legs with 2 setae basad. T1 nearly 3X wider than long, with 3 setae in a transverse
row near posterior margin. T2 nearly 1/3 wider than long, Dl, D2, SD1, and SD2
present. T3 nearly 2X as wide as long, 1 D seta, SD1, and SD2 present. Thoracic
spiracle on T2 adjacent to T1. T1 leg extending to posterior margin of A3. T2 leg
extending nearly to anterior margin of A6. Forewing extending slightly beyond T2 legs.
T2-A3 with dorsal setae on a ridge. A9 and 10 taper to a point, intersegmental division
less conspicuous than other abdominal segment. Abdominal spiracles on A2-7 round
with those on A8 reduced and inconspicuous. Dorsal pinacula on A1-8 with 2 setae on


June, 1990

Cassani et al.: Sphenarches anisodactylus in Florida

or near a prominent bispinose process; trispinose and largest on A3. A3-7 with a pair
of small light colored bifurcate processes between D setae; one branch extending an-
terad, one posterad, best developed on A4, vestigial with one branch on A7. Al-8 with
SD1 and SD2 present. A2-7 with L1 directed anterad, L2 directed posterad. SD and L
pinacula distinctly posterior to spiracle on A2-7. SD directly above spiracle on A8 and
L setae joined by a narrow ridge, L1 distinctly anterior to spiracle, and 4x length of
L2, L2 distinctly posterior to L1 or L2 spiracle. Cremaster and anterior midventral
part of A9 with dense covering of fine hook-tipped setae, variable in length.
ADULT. Adult males and females are 6.5 mm 0.4 mm and 6.3 0.4 mm in body
length respectively; the wingspan is 14.5 mm 0.6 mm for males and 15.0 mm 0.5
mm for females (n = 15). Genitalia and other adult characters are figured and described
in Matthews (1989).


The host plant, Thalia geniculata (Marantaceae), is a perennial herb arising from
thick rhizomes. Flowers occur along conspicuous zigzag (alternate) internodes on a
widely and loosely branched panicle. Of the approximately 400 species in this family,
all are exclusively tropical except T. geniculata. (Tarver et al. 1979). According to
Godfrey & Wooten (1979), T. geniculata occurs along the coastal plain of South Carolina
to Texas, Oklahoma and Missouri. Thalia geniculata is also reported from Africa and
the West Indies (Small 1933, Evans 1979). In Florida, T. geniculata is established
primarily in southern Florida and along the St. Johns River system (Tarver et al. 1979).
Populations of Thalia, along with natural wetland areas in Florida, are disappearing as
a result of development and altered hydrological cycles. If T. geniculata is the only host
plant for S. anisodactylus in Florida, then the distribution or existence of this moth
may parallel that of its host. The three T. geniculata populations studied in Lee County
occurred in a cypress swamp, roadside ditch, and a lowland pasture.
In South Florida, Thalia growth and abundance was closely synchronized with the
hydrological cycle. Seasonal regrowth begins in April or May usually the beginning of
the wet season. At the sites studied, flowering began during June or July and peaked
in October. December and January are relatively dry months in this area and the above
ground portions of the plant recede greatly at this time. By the end of January, the
study sites had become dry and little or no green growth was apparent until rains
initiated regrowth later in the season.
Our field observations indicated that seasonal occurrence of S. anisodactylus is
closely tied to flowering activity of Thalia. The earliest oviposition observed was in
July, 1979, when an estimated five percent of the plants were in flower. Eggs are laid
singly, primarily on the flower bracts. As many as 15-20 eggs per flower were not
Larvae were most abundant in August and by late September all life stages were
common on the flowers. Adults are inconspicuous and were occasionally observed rest-
ing on the leaves. In November 1979, 82 of 98 panicles observed at one site had at least
one life stage present. The pupa is generally attached by the cremaster to a silken pad
spun by the mature larva on the internodes or at the base of a bract if the flower has
not already fallen from the panicle.
By late December, only an occasional unemerged pupa could be found. The life stage
that overwinters is unknown, although a viable pupa was collected as late as 7 March
in Collier County and emerged on 13 March.
Based on our laboratory results, the duration of the immature stages is approxi-
mately 22 days and adults live 14-19 days (Table 1). As many as four or five generations
may occur from late July through December in South Florida. The duration of S.

Florida Entomologist 73(2)

June, 1990

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Cassani et al.: Sphenarches anisodactylus in Florida

anisodactylus immature stages is relatively short compared to the artichoke plume
moth, P. carduidactyla, which requires 80-110 days at outdoor temperatures in Califor-
nia (a more northern latitude than south Florida) (Lange 1941).
Mating occurs during the night and probably during predawn hours. Pairs in copula
may remain in an end to end position for several hours. Ovipositing females were never
directly observed but oviposition probably occurred at or before dawn since none were
observed ovipositing after 0700 hours or before 1800 hours. Lab reared females ovipo-
sited over a 2-6 day period with 1-22 eggs per female laid daily (minimum = 19,
maximum = 64). Lange (1941) found that the artichoke plume moth had a preoviposi-
tional period of 3-8 days with an average of 170 eggs laid per female. The relatively
long duration of oviposition probably contributes to the numerous overlapping genera-
tions of S. anisodactylus field populations in Florida. Numerous overlapping genera-
tions are also characteristic of the genus Platyptilia (Lange 1950).
The egg stage lasted 3 days (Table 1). Upon hatching, some larvae attempted to
chew the chorion of adjacent unhatched eggs, and later migrated to the petals or bored
into unopened portions of the flower. Rarely were more than two mature larvae found
feeding on a single flower in the field, indicating migration to other flowers or possible
mortality from unknown causes. Spiders are common predators on the panicle and could
account for the heavy losses of larvae that remain exposed on outer bud surfaces.
Larvae were observed feeding on all portions of the flower, especially the ovary. Larvae
have 4 instars, occurring over an 11 day period in the laboratory (Table 1). Mature
larvae move to the flower petiole (site of pupation), cease feeding, and become pale
green in color. The mean duration of the prepupal stage was one day (Table 1). The
pupal stage was approximately 7 days for both sexes (Table 1).
It is also interesting that Sphenarches anisodactylus is a pest of leguminous plants,
dicots, throughout much of the tropics but is known only from Thalia geniculata, a
monocot in Florida. In both cases, larvae prefer to feed in flowers and young pods. The
only other pterophorid known to feed on a monocot is, Platyptilia jezoensis from Japan,
which feeds on Alliumfistulosum L. (Amaryllidaceae) (Yano 1963). It is likely that S.
anisodactylus feed on Thalia in the tropics but this has not yet been reported. It is
also feasible that Sphenarches anisodactylus may use additional hosts in Florida. Pigeon
pea is not grown commercially in Florida but there may be small plantings for home
use among recent immigrants from the tropics. There are also at least two species of
Caperonia in Florida (Wunderlin 1982) which may serve as alternate hosts. Further
biological studies may explain the extreme polyphagy of this species. Perhaps different
biological races of S. anisodactylus occur that feed on different hosts. Secondary com-
pounds used by S. anisodactylus for host recognition may also be similar in these unre-
lated plants.


We thank Margo Duncan for the preparation of the illustrations. Mr. Paul Skelley
and Dr. Steven Passoa reviewed and improved the manuscript. Dr. Tomas Zoebisch
provided the Spanish abstract. Florida Agricultural Expreriment Station, Journal
Series No. R-00157.


ADAMCZEWSKI, S. 1951. On the systematics and origin of the generic group Oxyptilus
Zeller (Lep. Alucitidae). Bull. British Mus. (Nat. Hist.) Entomol. 1(5): 301-388,
pls. 9-20.

266 Florida Entomologist 73(2) June, 1990

BRUNER, S. C., L. C. SCARAMUZZA, AND A. R. OTERO. 1975. Catalogo de los
insects atacan a las plants economics de Cuba. 2nd Ed. Acad. Ciencas de
Cuba. 3.
COTES, E. C. 1891. Miscellaneous notes from the entomological section of the Indian
museum. Indian Museum Notes 2(1): 20-21,
EVANS, R. K. 1979. Guide to the wetland plants of submerged and transitional zone
lands. Florida Dept. Environ. Regulation. 294 pp.
FORBES, W. T. M. 1930. Insects of Puerto Rico and the Virgin Islands. Heterocera
or moths (excepting the Noctuidae, Geometridae and Pyralidae). Sci. Survey of
Puerto Rico and the Virgin Islands. 12(1): 1-171.
GODFREY, R. K., AND J. W. WOOTEN. 1979. Aquatic and wetland plants of the
southeastern United States, Monocotyledons. The University of Georgia Press,
Athens. 712 pp.
HEINRICH, C. 1916. On the taxonomic value of some larval characters in the Lepidopt-
era. Proc. Entomol. Soc. Washington 18: 154-164.
E. G. MUNROE, AND J. A. POWELL (eds.). 1983). Check list of the Lepidoptera
of America north of Mexico. E. W. Classey Limited and the Wedge Entomolog-
ical Research Foundation, London. 284 pp.
KIMBALL. C. P. 1965. The Lepidoptera of Florida an annotated checklist. Division of
Plant Industry, Florida Dept. of Agr., Gainesville, Florida. 363 pp.
LANGE, W. H. 1941. The antichoke plume moth and other pests injurious to the globe
artichoke. California Agr. Exp. Sta. Bull. 653: 1-71.
LANGE, W. H. 1950. Biology and systematics of plume moths of the genus Platyptilia
in California. Hilgardia 19: 561-668.
MATTHEWS, D. L. 1989. The plume moths of Florida (Lepidoptera: Pterophoridae).
MS Thesis, University of Florida, Gainesville, Florida. 346 p.
MCDUNNOUGH, J. 1927. Contribution toward a knowledge of our Canadian plume
moths. Trans. Royal Soc. Canada. Section V: 175-188.
MEYRICK, E. 1910. Lepidoptera Heterocera family Pterophoridae, pp. 1-23 in P.
Wytsman (ed.), Genera Insectorum. 100: 1-23.
MUNROE E. 1983. Pterophoridae, pp. 86-87 in R. W. Hodges (ed.), Checklist of the
Lepidoptera of America north of Mexico. E. W. Classey Limited and the Wedge
Entomological Research Foundation, London.
NEUNZIG, H. H. 1987. Pterophoridae, pp. 497-500 in F. W. Stehr (ed.), Immature
Insects. Kendall/Hunt. Dubuque, Iowa.
SMALL, J. K. 1933. Manual of the southeastern flora. Hafner Publishing Co., New
York. 1554 pp.
STANEK, V. J., B. TURNER (eds.). 1977. The illustrated encyclopedia of butterflies
and moths. English version. Octopus Books Limited-London. 352 pp.
STEHR, F. W. (ed.). 1987. Immature insects. Kendall/Hunt Publishing Co., Dubuque,
Iowa. 754 pp.
TARVER, D. P., S. A. RODGERS, M. J. MAHLER, AND R. L. LAZOR. 1979. Aquatic
and wetland plants in Florida. Florida Dept. Nat. Res., Tallahassee, Florida. 127
WALSINGHAM. T. 1897. Revision of the West-Indian microlepidoptera, with descrip-
tions of new species. Proc. Zool Soc. Lond. 1897: 56-62.
WUNDERLIN, R. P. 1982. Guide to the vascular plants of central Florida. Univ. of
South Florida, Univ. Presses of Florida, Tampa. 472 pp.
YANO, K. 1963. Taxonomic and biological studies of Pterophoridae of Japan (Lepidopt-
era). Pac. Insects 5(1): 65-209.

Schuster & Reyes-Castillo: Larval Passalidae 267


Universidad del Valle de Guatemala
Apartado 82
Guatemala, GUATEMALA

Institute de Ecologia
Apartado Postal 63
91000 Xalapa
Veracruz, MEXICO


The larvae of Didimus africanus (Percheron), D. haroldi Kuwert, D. alvarodoi (?)
Corella, D. parastictus (Imhoff), D. nachtigali Kuwert, Erionomus planiceps (Esch.),
E. pilosus Arrow, Taeniocerus pygmaeus (Kaup) and Comacupes basalis (Smith) are
described for the first time. Larvae of Leptaulax bicolor (F.), L. dentatus (F.) and
Aceraius grandis (Burm.) are redescribed and compared with previous descriptions
based on Indian specimens.
Passalid larval literature is summarized to date; a total of 120 species (29 Old World)
are now described.


Se described las larvas de Didimus africanus (Percheron), D. haroldi Kuwert, D.
alvaradoi(?) Corella, D. parastictus (Imhoff), D. nachtigali Kuwert, Erionomus plan-
iceps (Esch.), E. pilosus Arrow, Taeniocerus pygmaeus (Kaup) y Comacupes basalis
(Smith) por primera vez. Se redescriben las larvas de Leptaulax bicolor (F), L. dentatus
(F) y Aceraius grandis (Burm.), comparAndolas con descripciones previas basadas en
especimenes de la India. La literature sobre larvas de pasdlidos se resume hasta la
fecha; un total de 120 species, 29 del Viejo Mundo, son ahora conocidas.

This article summarizes the knowledge concerning passalid beetle larval taxonomy
and adds new larval descriptions. Schuster & Reyes-Castillo (1981) summarized the
literature to that date, a total of 22 species. They added 69 more species descriptions,
redescribed 6 species and provided a key to New World genera. Since then, Paulian &
Lumaret (1979) have described 3 species from Madagascar; Quintero & Reyes-Castillo
(1983) described 2 species of Oileus and redescribed the remaining 3 species in the
genus; Costa and da Fonseca (1986) have described 14 and redescribed 7 species from
Brazil with excellent drawings; Reyes-Castillo et al. (1987) described 2 Mexican species
of the new genus Xylopassaloides; and Schuster (1988) described Petrejoides reyesi
Schuster. This brings the total species described to 113, perhaps 1/6 of the world total.
Only 22 Old World species have been described. Here we describe 7 more Old World
species and redescribe 3 others. The methods and terminology (see Figs. 1 & 3) are the
same as in Schuster & Reyes-Castillo (1981). Collection locality and date follow the

'Research Associate, Florida State Collection of Arthropods, Dept. of Plant Industry, Gainesville, Florida.

268 Florida Entomologist 73(2) June, 1990

species name including the initials of which author collected the specimens. This is
followed by the number of individuals examined of each instar, with the range in head
widths (mm) for that instar.


Comacupes basalis (Smith) 1852 PHILIPPINE IS., Mindoro, S. of San Teodoro 18XII79
JCS 2III4.7-4.8
This species is known only from the Philippine Is. (Gravely 1914). The larva conforms
to the key characteristics for the subfamily (Schuster & Reyes Castillo 1981): re-
tinaculum present, antenna sub-bifurcate, trochanter-femoral stricture with setae. The
uncus of the lacinia is bifid. The primary setal pattern is quite complex (Fig. 1). The
labrum is bordered by many long setae. The frons has only short setae. Many short
hairs occur behind and below the antennae. The dorsal setal pattern is characterized by
short (0.25mm) primary setae. Up to 12 setae can occur in a transverse row on the
abdominal tergites. A single seta occurs on each lateral lobe below the level of the
spiracles on each abdominal segment, similar to those mentioned for Aulacocyclus eden-
tulus (MacLeay) (Schuster & Reyes-Castillo 1981). These we here call pleural seta (PS).
The 12 AR setae are all dorsal, as in A. edentulus (Schuster & Reyes-Castillo 1981).
The ventral abdomen is bare except for the abundant hairs on the raster. The anus is

Taeniocerus pygmaeus (Kaup) BRUNEI 22km S.W. Telisai (rd. to Labi) 20XI80 W.D.
Edmonds 44III2.0-2.3, 4111.5-1.6
This species occurs in peninsular Malaya, Sumatra and Borneo. The uncus of the
lacinia is entire. As in other Aulacocyclinae, a retinaculum is present, the antennae are
sub-bifurcate and the leg stricture possesses setae. Head width distributions were:
2.0-2.05: 6, 2.1-2.15: 27, 2.2-2.25: 10, 2.3: 1. The internal coxae have 2-5 setae. The frons
is bare except for short (<0.1mm) setae in lateral corners. One pair of strong, and 2
pairs of weaker, post-antennal setae on the head (HPA) are present. Sclerotized pro-
notal shields are reduced to sunken points 0.3mm diameter or absent. The thorax has
1 or 2 pleural setae at spiracle level as well as approximately 5 long PS setae on each
pleural bulge (Fig. 2). Other hairs to 0.5mm occur on each abdominal segment, especially
near TL setae. The basic pattern includes 2 pairs of TD setae, but frequently a third
seta is present mesally. Usually 8 (37/44) anal ring setae occur, 6 in the dorsal hemis-
phere (Fig. 2). Each ventral abdominal segment is crossed by 10-16 long (0.8mm) setae,
though somewhat shorter posteriorly. The raster has short setae (instars II and III are
quite similar). Most unusual is the difference in size of the spiracles. Measured vertically
in instar III, their dimensions on abdominal segments 1-8 are (mm): 0.25, 0.15, 0.1, 0.1,
0.1, 0.15, 0.2, 0.2.
This description agrees with that of T. bicuspis Kaup (Gravely 1919) with respect
to the presence of pleural thoracic setae, though differing in number. Gravely's brief
description indicates fewer dorsal setae, 6 anal ring setae, and lack of any other hairs!


Aceraius grandis (Burmeister) 1847. TAIWAN, Ku Kuan 3180 JCS 7III6.2-7.0, 6II4.7-
4.9, 213.3-3.5
This species occurs from India to Taiwan and the Sunda Is. Previous descriptions
of the larvae of this species refer to Indian specimens (Gravely 1916, Gardener 1935).
The Indian specimens seem to be quite similar to those from Taiwan; however, the
previous descriptions apparently refer only to the second and third instars.

Schuster & Reyes-Castillo: Larval Passalidae

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270 Florida Entomologist 73(2) June, 1990

The uncus of the lacinia is bifid. The primary setal pattern includes 1-3 long setae
in each lateral angle of the frons, 5-7 very long HPA setae in a vertical row, 3 pairs of
TM setae forming a transverse row on abdominal tergites 2-6, occasionally on tergites
1 or 7, and, in the third instar, occasionally on tergite 8. The anal ring has 18-20 setae
in all instars. Usually a pair of short AV9 setae is present. The anus is a single trans-
verse slit. A hair pile occurs over most of the body. The raster is quite hairy. The first
instar larva differs strikingly from the others (Fig. 3). It has a more complete setal
pattern in that there are 3 pairs of TM setae on abdominal segments 1-9 as well as on
each thoracic segment (1 or 2 individual setae may be lacking in the latter site). In
addition, there are a single pair of TL and 2 pairs (the anterior seta shorter) of MSL
and MTL setae. The anterior corner of the pronotum has 7-9 PSL setae, with the
weakest ones located more mesally. Usually, 3 pair of PD, MSD, and MTD setae are
present, often with a short extra pair slightly anterior to the outer MSD and MTD setae.

Leptaulax dentatus (Fabr.) 1792 PHILIPPINE IS., Luzon, Los Bafios, University of the
Philippines 14XII79 JCS 5III4.3-4.6
This species occurs from India to the Philippines, Sunda Is., Moluccas, New Guinea
and Australia. The larva was originally very briefly described by Gravely (1916) from
4 Indian specimens. He mentions only the distribution of some hairs and setae. In our
specimens the uncus of the lacinia is entire. The mandibles are without a retinaculum,
as in all other Passalinae we describe here. The frons and HPA setae are all short
(<0.2mm), not clavate. The primary setal pattern (Fig. 4) includes only a single pair of
TM setae on tergites 1-7 [sometimes lacking on tergite 7 or present as far as tergite 8,
according to Gravely (1916)]. The anal ring has 12 setae (13 in one specimen). No AV9
setae are present. The mid-ventral raster is bare; a few short (<0.2mm) hairs are
present laterally. Hairs are scarce on the body. The coxae have one (rarely 2) internal
setae each. The anus is T shaped.

Leptaulax bicolor (Fabr.) 1801 PHILIPPINE IS., Mindoro, S. of San Teodoro
18XII79JCS 1III3.5, 3II2.2-2.3
This species also is found from India to the Philippines, Sunda Is., Moluccas, New
Guinea and Australia. It was originally described by Candeze (1861) with redescriptions
by Gravely (1916) and Gardner (1935) of Indian specimens. Gravely (1916) divided the
species into 2 varieties: L. bicolor s. str. and vicinus (Hope), the latter from the Anda-
man Is. Gravely apparently worked with all instars, Gardner with only third instars.
The uncus of the lacinia is bifid. According to Gravely (1916), the first instar of L.
bicolor s. str. has a single long HPA seta, lacking in other instars and reduced in variety
vicinus. Our specimens have 2 stout, short (<0.1mm) clavate setae as well as various
thinner clavate setae behind the antennae. He implied a lack of dorsal thoracic setae
early in development with ". . in a specimen little over 15mm . a single pair of
rather small dorsal hairs has appeared on each thoracic segment." Dorsal thoracic setae
were also mentioned by Gardner (1935); however, none are present on our specimens.
We agree in the presence of 3 PSL setae (sometimes 2 PSL according to Gardner), as
well as the basic primary setal pattern (Fig. 5) of 2 pair of TM setae on tergites 1-9
plus a single pair of short, dorsally directed pleural setae on the lateral lobes just below
the spiracle line on abdominal segments 1-8 or 1-9. This latter seta in our specimens is
clavate. The anal ring has 14 setae. The raster is bare. On our specimens there are no
other setae or hairs on the body larger than 0.1mm except on the legs, prosternum and
mouthparts. There are 2 internal coxal setae. Gravely (1916) and Gardner (1935), how-
ever, mentioned what is apparently a pair of AV9 setae. Our specimens are smaller
than those of Gravely and Gardner. Gravely (1916) mentioned associated adults 29mm
long, whereas our adults measure approximately 18mm. L. bicolor is geographically
very wide ranging; therefore, without examination of further specimens from different

Schuster & Reyes-Castillo: Larval Passalidae 271

areas, it is impossible to determine if we are dealing with more than 1 species or a
highly variable single species.

Didimus Kaup

Our 5 species have the uncus of the lacinia entire, frons, HPA and most thoracic
setae very short (<0.1mm). A basic setal pattern seems characteristic of the genus (Fig.
6). The trochanter-femoral constriction is bare except for 1 dorsal hair. The tergum has
scattered secondary setae <0.1mm long. The raster and ventral abdomen are bare; the
prosternum has a few hairs. The anus is T- or Y-shaped. The species differ in the
number of internal coxal setae, head capsule width and some setae. The basic anal ring
setal number is 12. The following key may be used to separate our species:


1 Internal coxa with 3 setae, 4 small setae between major TM setae, TM 1-6
setae long .............................................................................. D africanus
1' Internal coxa with <3 setae, no small setae between major TM setae, TM 1-9
setae long .............................................................................................. 2
2 Internal coxa with 1 seta, meso- and metanota with 1 pair MSL and MTL
setae; instar II head width 2.1- 2.2mm ....... ... ................ D. nachtigali
2' Internal coxa with 2 setae, meso- and metanota with 2 pairs MSL and MTL
setae; instar II head width >2.4mm .................................................... 3
3 Long PD, MSD, and MTD setae present, instar II head width 2.7mm, instar
III head width >4.2mm ............................................................... D. haroldi
3' Long PD, MSD, and MTD setae absent, instar II head width <2.7mm, instar
III head width <4.2mm .................................................. ................... 4
4 Pronotum with 5 pairs of short PSL setae ............................... D. parastictus
4' Pronotum with 0-3 pairs of short PSL setae ........................... D. alvaradoi (?)

Didimus africanus (Percheron, 1844) IVORY COAST, Tai National Pk. 2VIII80 PRC
This species occurs in west and east tropical Africa. The internal coxal area has 3
setae. The diameter of PSL setae becomes smaller mesally, with at least 3 definite thick
setae near the external anterior corner of the shield. The only other notal setae are a
pair of MSL and MTL setae. Long TM setae are present on abdominal tergites 1-6,
with short truncated setae on the other segments. Internal to the TM setae are 2 pair
of very short, pointed, thinner (but thicker than other body hairs) setae. A pair of short,
truncated TL setae occurs on each abdominal tergite.

Didimus parastictus (Imhoff, 1843) IVORY COAST, Tai National Pk. 2VIII80 PRC
7III3.6-3.8, 2112.6
This species occurs in west tropical Africa. The internal coxa has 2 setae. The PSL
setae resemble those of D. africanus with at least 5 setae thicker than the others in
each anterior lateral corner of the shield. Two pairs of MSL and MTL setae are also
present. A pair of TM setae is usually complete on each abdominal tergite with TL setae
on tergite 9 only. Seven larvae had 12 AR setae, 1 had 11, 1 had 10. Second and third
instars were essentially the same; both possessed metanotal bars. One red egg in early
stage of development was found.

D. nachtigali Kuwert (1891) IVORY COAST, Tai Nat. Pk. 30VII80 PRC 2112.2, 111.4;
5VIII80 PRC 1III3.2, 1112.2

Florida Entomologist 73(2)

This species occurs in tropical east and west Africa. The internal coxa has only 1
seta. MTD setae are lacking, this being the principal difference from what is probably
the basic pattern for the genus. All instars had the same setal pattern. Metanotal bars
were present only on instar I. Two red-brown eggs 1.9 x 1.5mm were found.

D. haroldi Kuwert 1898 IVORY COAST, Tai National Pk., VIII80 PRC 3III4.3-4.4,
This species occurs in tropical east and west Africa. The internal coxa has 2 setae.
It differs from the basic setal pattern in lacking PSL setae, possessing an extra pair
each of MSL and MTL setae and all lateral setae are reduced to stubs except TL9.
Instar II lacked MTD setae, possessed metanotal bars, but otherwise was the same as
instar III.

D. alvaradoi (?) Corella IVORY COAST., Tai National Pk. 30VII80 PRC 1112.5, 211.7-1.8
This species is known from Spanish Guinea. The internal coxa has 2 setae. It differs
from the basic setal pattern in lacking: (1) PSL setae in instar II; (2) PD, MSD, and
MTD setae in instars I and II (except for remnants of MTD in instar II); (3) TSL 1-8
setae in instar II (reduced to stubs in instar I). It also has 2 pair of stubby MSL and
MTL setae. Metanotal bars are present in instar I.

Erionomus Kaup

Our 2 species have the uncus of the lacinia entire. Frons setae are <0.01mm long.
Two to 4 HPA setae longer than the antenna are oriented in a vertical row; smaller
thick setae are also present. The basic setal pattern is that of Fig. 7. Raster and venter
are bare, prosternal hairs are <0.01mm long. The anus is Y or T shaped.

E. planiceps (Esch.) IVORY COAST, Tai National Pk 5VIII80 PRC 2III7.0, 213.1,3.2;
31VII80 PRC 1114.5, 113.1
This species occurs in east, equatorial, and west Africa. It has 3-4 long HPA setae,
3 or 4 internal coxal setae, and scattered secondary setae (<0.2mm) dorsally. All instars
have the basic dorsal setal pattern, with occasional additions of another pair of internal
TM, PD and MSD, as well as an occasional extra pair of MSL setae, especially in later
instars. Metanotal bars are present in instars I and II. The anal ring contains 18 setae
(17-19 in instar I).

E. pilosus Arrow IVORY COAST, Tai National Pk. 3VIII80 PRC 112.6 1II (molting)
This species occurs in west and equatorial Africa. The instar II head width is not
given because it was in ecdysis when it died. The specimens have 2-3 long HPA setae,
2-3 internal coxal setae, but no secondary setae dorsally. They have the basic
Erionomus dorsal setal plan. The anal ring contains 16 setae. A brown egg 3.1 x 3.6mm
with embryo undeveloped was found.


JCS thanks Tom Shih, Taichung University, Toto Mapa, John Somebang, Isabel
Walter, and the Philippine Match Corp. for incredible hospitality and aid in collecting
passalids; and the Universidad del Valle de Guatemala for support. PRC thanks Prof.
Lamotte, Ecol'e Normale Superieure, Paris, and Y. Cambefort for facilitating field
work in the Ivory Coast; W. D. Edmonds for larvae of T. pygmaeus; Gonzalo Halffter
del Instituto de Ecologia, Mexico, F. di Castri, UNESCO MAB program, CONACYT-
Mexico and the Institute of Ecology, Ivory Co., for support. We thank Michael C.
Thomas and an anonymous reviewer for comments on the manuscript.


June, 1990

Wirth & Linley: Dasyhelea chani 273

Contribucion #15 al proyecto PCECCNA-040451 apoyado por el Consejo Nacional
de Ciencia y Tecnologia (M6xico) y desarrollado parcialmente en el Departamento de
Biosistematica de Insectos, Instituto de Ecologia.


CANDEZE, E. 1861. Histoire des M6tamorphoses de quelques Coleopteres exotiques.
Mem. Soc. R. Sci. Liege 16: 343-344.
COSTA, C., AND C. R. V. DA FONSECA. 1986. Larvae of Neotropical Coleoptera.
XIII. Passalidae, Passalinae. Revta. bras. Ent. 30(1): 57-78.
GARDNER, J. C. M. 1935. Immature stages of Indian Coleoptera (16), Scarabaeoidea.
Indian Forest Rec. (New Ser.) Ent. 1(1): 1-33.
GRAVELY, F. H. 1914. An account of the Oriental Passalidae (Coleoptera). Mem.
Indian Mus. 3: 177-359.
1916. XIV Some lignicolous beetle-larvae from India and Borneo. Indian Mus.
Rec. 12: 137-175.
1919. XVII Descriptions of Indian beetle larvae III. Indian Mus. Rec. 16:
PAULIAN, R., AND J. P. LUMARET. 1979. Famille des Passalidae. Faune de Madagas-
car 50: 11-50.
QUINTERO, G., AND P. REYES-CASTILLO. 1983. Monografia del g6nero Oileus Kaup
(Coleoptera, Scarabaeoidea, Passalidae). Folia Entom. Mex. 57: 1-50.
REYES-CASTILLO, P., C. R. V. DA FONSECA, C. CASTILLO. 1987. Descripcion de
un nuevo g6nero mesoamericano de Passalidae (Coleoptera: Lamellicornia). Folia
Entomol. Mex. 73: 47-67.
SCHUSTER, J. 1988. A description of Petrejoides reyesi sp. nov. (Coleoptera, Pas-
salidae) from Honduras. Coleopterists Bull. 42(4): 305-309.
AND P. REYES-CASTILLO. 1981. New World genera of Passalidae (Coleopt-
era): a revision of larvae. An. Esc. nac. Cienc. biol., Mex. 25: 79-116.


Research Associate, Florida State Collection of Arthropods
1304 NW 94th St., Gainesville, FL 32606

Florida Medical Entomology Laboratory, Institute of Food & Agricultural Sciences
University of Florida, 200 9th St. S.E., Vero Beach, FL 32962


Dasyhelea chani new species (Diptera: Ceratopogonidae), whose immature stages
are found on leaves of the water lettuce, Pistia stratiotes L., is described in the pupal
and adult stages.

Florida Entomologist 73(2)


Se describe en el estado pupal y adulto a Dashyelea chani nueva especie (Diptera:
Ceratopogonidae), cuyas estapas inmaduras se encuentran en hojas de la lechuga de
agua, Pistia stratiotes L.

Biting midges of the genus Dasyhelea Kieffer are common and widespread, found
in all regions of the world and in a wide variety of habitats. The larvae are aquatic or
semiaquatic, requiring at least a thin film of water in which to live. They are unable to
swim, so they move by climbing or hitching their way along, using their mouthparts
and posterior hooks. Their preferred habitats are wet moss or algae along shores of
streams, lakes, ponds, puddles and other bodies of water, or in wet rotting materials
such as sap oozing from trees, wet bark and tree holes. Some species are found in
unusual habitats such as rock pools, thermal water in hot springs, or water with high
mineral or salt content. They have successfully invaded the tidal zone along seashores,
where many species breed on algae-covered rocks or in algae growing on mud exposed
to tidal action in salt marshes. Most larvae spin tubular cases in the last instar.
Adult Dasyhelea midges may be found in habitats near the breeding sites. Some
species are found among shrubs, on plants near water, or at flowers. Little is known
about the feeding habits of adults. As far as known, adults feed on honeydew and sweet
secretions of plants, or they visit flowers for nectar. Blood-sucking habits are not known
to have been developed in this genus. Some species are important pollinators of cacao
and other tropical tree crops.
Approximately 500 species of Dasyhelea have been described worldwide, of which
only 42 are known from North America. Waugh & Wirth (1976) listed only 18 species
in the genus from the Eastern United States north of Florida, and Wilkening et al.
(1985) recorded 15 species for the state of Florida. Thus only a small fraction of the
existing North American Dasyhelea species have been described, and there are
thousands of specimens in the collection of the National Museum of Natural History of
the Smithsonian Institution in Washington awaiting study and description.


The material forming the basis of the present study was collected by Dr. Kai Lok
Chan during studies on the biology of the ceratopogonid midges associated with water
lettuce plant (Pistia stratiotes L.) at Chinese Farm, an abandoned aquaculture project
adjacent to Old Dixie Highway, about 3 miles south of the Florida Medical Entomology
Laboratory at Vero Beach, Florida.
Our taxonomic material was killed by immersion in hot water and preserved in 70%
ethanol. Adults were dissected and mounted on microscope slides in phenol balsam
according to the technique of Wirth & Marston (1968). Terminology used in description
of the adult midges follows that of Waugh & Wirth (1976) and Downes & Wirth (1981).
Terminology for the pupal description follows that used for Culicoides by Blanton &
Wirth (1979). The holotype and allotype and a portion of the paratypes are deposited
in the National Museum of Natural History in Washington; additional paratypes are in
the collections of the authors and the Florida State Collection of Arthropods in Gaines-
ville, Florida.

Dasyhelea chani Wirth and Linley, new species (Fig. 1-14)

Diagnosis. A small brownish species with yellowish or pale brown scutellum, dark
halters and pale yellowish to brownish legs; wing whitish with sparse, coarse, sharp,


June, 1990

Wirth & Linley: Dasyhelea chani 275

,------ ----,-


7 8

Fig. 1-8. Dasyhelea chani; 1, 6, 8, male; 2-5, 7, female: 1, 2, antenna; 3, palpus; 4,
wing; 5, spermatheca; 6, parameres; 7, female distal abdominal segments, ventral view;
8, male genitalia, parameres omitted.
spinelike macrotrichia in linear arrangement. Pupa pale yellowish; abdominal tubercles
small; respiratory horn tapering to sharp point apically, with slitlike spiracular openings
on distal portion; last segment with two pairs of long, slender, spinelike terminal proces-
Female Holotype. Wing length 0.90 mm, breadth 0.42 mm; costal ratio 0.48.
Head. Brown; antenna dark brown, palpus paler at base. Antenna (Fig. 2) with

Florida Entomologist 73(2)


Fig. 9-14. Dasyhelea chani; 9, operculum; 10, prothoracic respiratory horn; 11, ab-
dominal segment 8, dorsal view; 12, abdominal segment 4, dorsal view; 13 same, ventral
view; 14, last abdominal segment of female pupa, dorsal view.

June, 1990

Wirth & Linley: Dasyhelea chani 277

lengths of flagellar segments (in microns) 39-26-29-29-31-31-31-31-43-45-45-43-93; anten-
nal ratio (elongated segments 11-15/shorter segments 3-10) 1.10; proximal segments
globular, slightly elongated toward 10; 11-13 about twice as long as basal breadth,
slightly tapering; 15 tapering to slender tip; all segments with definite sculpturing.
Palpus (Fig. 3) short and slender; segments 1 and 2 incompletely divided, more distinct
than normal in the genus; lengths of segments 1 + 2 to 5 (in microns) 36-36-29-44; 3rd
segment globular to slightly longer than broad; palpal ratio 1.67 in holotype.
Thorax. Dark brown, scutellum paler brown. Legs stramineous, knee spots slightly
darkened, tarsi slightly darker distally; provided sparsely with rather long dark setae;
lengths of segments of hind leg from femur distad (in microns) 360-340-232-90-76-51-51.
Wing (Fig. 4) with membrane whitish due to well-developed microtrichia; macrotrichia
coarse, long, spinelike, and sparse, arranged in lines paralleling veins; 1st radial cell
obliterated, 2nd slitlike, end of costa oblique. Halter brownish.
Abdomen. Dark brown; terga deeply pigmented blackish with pale punctures at the
bases of the sparse, rather coarse setae; pleura appearing blackish due to dense coarse
spicules arranged in close-set, longitudinal rows. Subgenital plate (Fig. 7) a stout, bi-ar-
cuate, transverse band bearing a short, broad, slightly bilobed anteromedian projection.
Spermatheca (Fig. 5) one; large, moderately pigmented, subspherical to slightly ovoid
with a short, slender, slightly oblique neck; measuring 0.072 by 0.061 mm with neck
0.007 mm long.
Male Allotype. Wing length 1.04 mm, breadth 0.38 mm; costal ratio 0.45.
Similar to the female with the usual sexual differences. Thorax darker brown than
in female, scutellum paler yellowish; legs with fore and hind femora and tibiae more
brownish in midportions. Antenna with lengths of flagellar segments (in microns) 61-36-
36-35-33-35-38-38-41-80-73-64-108; antennal ratio (12-15/3-11) 0.91; segments deeply
sculptured; plume dense, blackish, with long verticils reaching midlength of segment
15. Wing as in female but narrower, macrotrichia sparser. Lengths of segments of hind
leg from femur distad (in microns) 414-370-280-111-90-58-58.
Genitalia (Fig. 8). Ninth segment slightly broader than long; sternum slightly pro-
duced caudomesad abutting base of aedeagus with a submedian pair of strongly
sclerotized, low prominences; tergum rounded caudad with a small submedian pair of
beadlike apicolateral processes, each bearing a short seta. Basistyle about twice as long
as broad, with a low setose ventromesal lobe distally, and a strongly sclerotized hooklike
process on midway mesal margin as in species of the grisea Group; dististyle about as
long as basistyle, only slightly curved, and moderately tapering to slender tip. Aedeagus
complicated; with a narrow, transverse, slightly arcuate, strongly sclerotized, basal bar
with a low, bluntly pointed, caudomedian expansion; from which posteriorly arise caudo-
ventrally a pair of short winglike plates each with posterior ends divided in two sharp-
pointed unequal lobes, from the outer lobe of which a ridged thickening curves arcuate
anterolaterall and dorsally toward ends of basal bar; these arcuate sclerites forming a
strong ventromesal support for a second pair of more dorsally and laterally located
plates which are blunt-pointed caudally and lightly sclerotized, and form the support
for a longitudinally wrinkled, hyaline envelope for the distal part of the male genital
duct. Parameres (Fig. 6) strongly asymmetrical as usual in the genus; basal apodemes
forming nearly straight, moderately broad bands; caudomedian sclerite slightly sinuate
and slightly swollen proximally in ventral view, evenly curving and tapering in lateral
view, with slender, tapering, distal process directed ventrad apically; an area of numer-
ous setae separates the posteriorly directed portion from the more slender, ventrally
directed, distal portion.

278 Florida Entomologist 73(2) June, 1990

Pupa. Length 2.9 mm. Exuviae pale yellowish throughout, last segment amber col-
ored; respiratory horn amber with hyaline tip and pale margin at spiracular openings.
Devoid of strong setae; integument smooth except for fine shagreening on abdominal
segments, coarser on posterior margins of segments and over all of last segment. Oper-
culum (Fig. 9) rounded ventrally, slightly bilobed dorsally, with a pair of small sharp
spinules laterally at widest portion; no setae or sculpture visible. Respiratory horn (Fig.
10) of unusual structure, relatively stout with parallel margins on proximal half, then
tapering and curving to a sharp distal point; about 16 inconspicuous slitlike spiracular
openings in a loosely arranged row on distal 0.4 of curved dorsal margin and distal 0.2
of ventral margin; internal tracheal chamber divided in two portions, the distal portion
bearing tubules leading to the spiracular openings, and proximal portion on proximal
third of horn with fine transverse annulations Abdomen (Fig. 12-13) with 1st segment
reduced and 2nd elongated as usual in the genus; segments 3-7 each with 9 pairs of small
bluntly-pointed posteromarginal tubercles, each with a minute seta, aligned in a nearly
straight ring around posterior margin of segment, the ring broadly interrupted on
dorsal and ventral midlines of segment, the two lateromost pairs of tubercles enlarged,
somewhat thornlike; segment 8 (Fig. 11) with only 5 pairs of such tubercles. Posterior
segment (Fig. 14) with unusual modification for the genus, somewhat similar to that of
Dasyhelea traverae Thomsen as figured by Waugh & Wirth (1976); instead of the usual
broad and flattened terminal processes, these are much elongated in long, slender,
minutely fringed, yellowish spines 0.20 mm long (or 1.25 as long as segment); at the
base of each terminal spine dorsally arises a much shorter, darker, minutely fringed
spine only 0.053 mm long, between these spines arises a minute hyaline seta about 0.005
mm long.
Larva. Length 4-5 mm when mature. Pale creamy white; head concolorous. Head
capsule 0.324 mm long by 0.200 mm wide; mouthparts and pharyngeal apparatus not
studied. Head and body bare of setae. Last segment with 2 dorsal pairs and 4 ventral
pairs of slender, inconspicuous hooks with some microscopic spinules interposed be-
tween the groups of hooks.
Types. Holotype female, allotype male; paratypes, 9 females, 8 males. FLORIDA,
St. Lucie Co., Ft. Pierce, Chinese Farm, 24 July 1987, K. L. Chan, reared from Pistia
stratiotes L. Other material: 12 pupal exuviae, 10 larvae, same data.


We dedicate this species with pleasure to Dr. Kai Lok Chan of the National Univer-
sity of Singapore, who, spending a sabbatical leave working at the Florida Medical
Entomology Laboratory, reared a most unusual variety of ceratopogonid midges from
leaves of water lettuce. Ecological studies on the relation of this species to Pistia and
its fauna will be reported separately by Dr. Chan and the junior author.
Dasyhelea chani is provisionally placed in the grisea Species Group on the basis of
its reticulated antenna with tapering, elongate, slender, last antennal segment; subgen-
ital plate with well-developed anteromedian lobe; male basistyle with well-developed
hooklike process on mesal margin; male aedeagus with several pairs of caudomedian
processes; and parameres asymmetrical with elongate caudomedian sclerite. The un-
usual development of the distally pointed respiratory horns and the long spinelike caudal
processes of the pupa are found in another species of the grisea Group, Dasyhelea
traverae Thomsen. Otherwise, D. traverae does not seem to be closely related, with
much different subgenital plate, male genitalia and wing, while the pupa differs in
bearing spiracular openings on both edges of the respiratory horn, and lacking the 2nd
short pair of spines on the last abdominal segment (Waugh & Wirth 1976). The sharp-
pointed, spinelike macrotrichia on the wing of D. chani are a good quick recognition

Wirth & Linley: Dasyhelea chani

feature, but this character is shared with D. spiniforma Waugh and Wirth, another
unrelated and otherwise dissimilar North American species.
A third Dasyhelea species (but unnamed) with a pupa similar to that of D. chani
was described and figured in the pupal stage by Mayer (1934) as a "Holoconops sp.".
The pupa was collected by Feuerborn during the German Limnological Sunda Expedi-
tion in Sumatra, where it was found among Sphagnum in a large freshwater pool.
The unusual modification of the pupal respiratory horn and apical abdominal segment
begs some explanation of its function. In this regard attention should be called to the
remarkable pupa of Stilobezzia poikiloptera Ingram and Macfie described by Ingram
and Macfie (1922) from specimens reared from Pistia stratiotes at Accra, West Africa.
While possessing the slender, sharp-pointed respiratory horn and long spinelike caudal
processes, the pupa of S. poikiloptera is otherwise quite different, belonging to another
subfamily of biting midges, the Ceratopogoninae. It would seem that such a parallelism
must serve a real biological function, such possibly as using these sharp processes as
spurs to aid in climbing or moving around on the plants.
Another question arises, which cannot be answered until the Dasyhelea fauna of the
tropics and subtropics worldwide has been more thoroughly sampled and studied. At
present our knowledge of the taxonomy and biology of this genus is fragmentary, nearly
the least-studied in the entire family of biting midges. Has Dasyhelea chani been
brought into Florida along with the introduction of its host plant, the water lettuce? Or
is it a rare and previously unknown native Florida species that has taken the opportunity
to adapt to an immigrant host that becomes locally abundant and dominant in its ecolog-
ical niche?


This paper is Institute of Food and Agricultural Sciences, University of Florida
Experiment Stations Journal Series No. 10018.


BLANTON, F. S., AND W. W. WIRTH. 1979. The sand flies (Culicoides) of Florida.
(Diptera: Ceratopogonidae). Arthropods of Florida and Neighboring Land Areas
10: 1-204.
DOWNES, J. A., AND W. W. WIRTH. 1981. Ceratopogonidae, p. 393-421 in J. F.
McAlpine et al. (eds.), Manual of Nearctic Diptera, Vol. 1. Monograph No. 27,
Biosystematics Research Institute, Ottawa, Ontario.
INGRAM, A., AND J. W. S. MACFIE. 1922. West African Ceratopogoninae Part II.
Ann. Trop. Med. Parasit. 16: 243-282.
MAYER, K. 1934. Ceratopogoniden-Metamorphosen (C. Inermediae und C. Ver-
miformes) der Deutschen Limnologischen Sunda-Expedition. Arch. f. Hydrobiol.
Supply. Bd. 13: 166-202, 6 plates.
WAUGH, W. T., AND W. W. WIRTH. 1976. A revision of the genus Dasyhelea Kieffer
of the eastern United States north of Florida (Diptera: Ceratopogonidae). Ann.
Ent. Soc. america 69: 219-247.
WILKENING, A. J., D. L. KLINE, AND W. W. WIRTH. 1985. An annotated checklist
of the Ceratopogonidae (Diptera) of Florida with a new synbonymy. Florida Ent.
68: 511-537.
WIRTH, W. W., AND N. MARSTON. 1968. A method for mounting insects on micro-
scope slides in Canada balsam. Ann. Ent. Soc. America 61: 783-784.

Florida Entomologist 73(2)


The Land, EPCOT Center, Lake Buena Vista, Florida 32830


Measurements of either the length of mouth hooks or the cephalopharyngeal skeleton
showed clear separation between larval instars of Liriomyza sativae Blanchard. Mouth
hooks of first instar larvae also differed morphologically from those of later instars.
Larval instars were distinguished under a dissecting microscope at 200x magnification,
enabling rapid separation of freshly-collected specimens.
The growth ratio of cephalopharyngeal skeletons between first and second instar
larvae was higher (1.75) than between second and third instar larvae (1.56). Growth
ratios calculated from published reports on other Liriomyza species were typically very
similar to those of L. sativae.


Medidas del largo de los ganchos bucales o del esqueleto cefalofaringeal demostr6
una clara separaci6n entire los estadios larvales de Liriomyza sativa Blanchard. Los
ganchos bucales tambi6n fueron distintos morfologicamente de aquellos en los estadios
siguientes. Los estadios larvales se separaron bajo un microscopio de disecci6n a una
magnificaci6n de 200X, lo que permiti6 separar rdpidamente los especimen reci6n colec-
La taza de crecimiento de los esqueletos cefalofaringeales entire el primer y segundo
estadio, fue mayor (1.75) que entire el segundo o tercer estadio larval (1.56). La taza de
crecimiento calculada en reports publicados sobre otras species de Liriomyza fue
tipicamente muy similar a aquellos de L. sativae.

Liriomyza leafminers cause extensive damage to a variety of economically important
vegetable and ornamental crops (Parrella & Keil 1984, Minkenberg & van Lenteren
1986). Consequently, they are major targets of both chemical (Parrella et al. 1982,
Mason et al. 1987, Leibee 1988) and biological control programs (Frijters et al. 1986,
Westerman & Minkenberg 1986, Parrella et al. 1987). These programs would likely
benefit from more detailed physiological or ecological studies conducted with particular
larval instars. Larval instars of cyclorrhaphous Diptera have shown differences in their
ability to escape parasitism and encapsulate parasitoid eggs (van Alphen & Drijver
1982), and in their susceptibility to pesticides (Parrella et al. 1982).
Larval instars of cyclorraphous Diptera are generally distinguished on the basis of
the size and/or morphology of the cephalopharyngeal skeleton, which is characteristic
in this group (Teskey 1981). The cephalopharyngeal skeleton consists of the mouth
hooks and the hypopharyngeal and tentoropharyngeal sclerites (Teskey 1981). The size
and/or morphology of mouth hooks (Bodenstein 1950, Kamali & Schulz 1973, Robinson
& Foote 1978, Lawrence 1979), and/or the length of the entire cephalopharyngeal skele-
ton (Beri 1974, Ipe 1974, Hendrickson & Barth 1978, Lawrence 1979) have been used
to distinguish larval instars.
In this study, larval instars of Liriomyza sativae Blanchard were clearly distin-
guished using measurements of mouth hook or cephalopharyngeal skeleton length on
slide-mounted larvae using a compound microscope. The measurements were used also


June, 1990

Petitt: Distinguishing Larval Instars of Liriomyza sativae 281

to develop a method to distinguish instars using a dissecting microscope equipped with
an ocular micrometer. Being able to conduct instar determination under a dissecting
microscope is desirable because it considerably reduces processing time for samples
containing many larvae.
Growth ratios of L. sativae mouth hooks and cephalopharyngeal skeletons were
compared both across instars and with other Liriomyza species. Constant growth ratios
were expected within a species according to the Brooks-Dyar rule of geometric growth
(Hutchinson & Tongring 1984, Daly 1985).


Experimental Procedure

Bush lima beans (Phaseolus limensis var. limenanus Bailey 'Henderson') were
grown in a greenhouse in 15 cm diameter pots at two plants per pot in a steam-pas-
teurized peat-vermiculite mixture (Speedling Inc., Sun City, FL) and watered with a
modified Hoaland solution (Petitt 1988). Plants 10-14 d old were placed for 6 h in a 1.9
x 1.1 x 0.6 m cage containing a colony of adult L. sativae. Thereafter, plants were held
in environmental chambers at 20 or 25 + 1IC and a 14 h photophase beginning at 0700
Larvae were collected from primary leaves every 6 and 12 h at 25 and 200C, respec-
tively, beginning the third day after oviposition and preserved in a 70% ethanol:water
mixture. Larvae were cleared in a 10% solution of potassium hydroxide in 70% ethanol
for 2-12 h, depending on larval size. The lengths of mouth hooks and cephalopharyngeal
skeletons (hypopharyngeal and tentoropharyngeal sclerites only) of 560 larvae were
measured on a compound microscope at 400x using an ocular micrometer (Fig. 2). Meas-
urements were made to the nearest micrometer unit which was equal to 2.5pm. Length
differences between the smallest mouth hook or cephalopharyngeal skeleton measured
for an instar and the largest of the preceding instar were used to determine the magnifi-
cation necessary for distinguishing instars using a dissecting microscope. Two hundred
larvae, some of which were from each stadium, were classified using a dissecting micro-
scope with an ocular micrometer to test the method and to determine the time required
for sample processing.

Data Analysis

The natural logarithms of the length of mouth hooks and the cephalopharyngeal
skeleton were regressed against presumed instar number (independent variable) to
confirm that no instars were missed (Daly 1985). Growth ratios were calculated by
dividing the size of a structure in an instar by the size of the equivalent structure in
the preceding instar.

Voucher Specimens

Specimens of adult L. sativae were sent to K. A. Spencer (Univ. of Exeter, Exwell
Farm, Bray Shop, Callington, PL17, 8QJ, Cornwall, United Kingdom) and their correct
identification was verified. Voucher specimens were deposited in the Florida State
Collection of Arthropods in Gainesville, Florida.


Clear separation among the three larval instars of L. sativae occurred using either
mouth hook or cephalopharyngeal skeleton length (Fig. 1; Table 1). No abrupt deviation

Florida Entomologist 73(2)

June, 1990


E 07
. 06





.04 .06 .08 .1 .12 .14 .16 .18
Length of Cephalopharyngeal

.2 .22 .24


Fig. 1. Relationship between lengths of mouth hooks and cephalopharyngeal skeleton
for first, second, and third instar L. sativae larvae.

occurred when presumed instar number was regressed with the natural logarithms of
mouth hook (r = 0.99) and cephalopharyngeal skeleton (r=0.99) length, indicating that
no instar was missed (Daly 1985). The mean cephalopharyngeal skeleton lengths of first,
second, and third instar L. sativae (0.09, 0.15, and 0.23 mm, respectively) were very
similar to the mouth hook lengths reported by Oatman & Michelbacher (1958) for L.
pictella Thomson. These authors apparently used the term "mouth hooks" to describe
the cephalopharyngeal skeleton. L. pictella of Oatman & Michelbacher (1958) actually
may have been L. sativae but the absence of voucher specimens makes positive identifi-
cation impossible (see Spencer 1981).
The mouth hooks of second and third instar larvae are morphologically similar, but
the first instar is recognizably different (Fig. 2). The second and third instar mouth
hooks were less than one-half as wide as long (including teeth) in side view, and had
two distinct pairs of alternating, ventrally curved, terminally-pointed teeth. The right
side of third instar mouth hooks was longer than the left in front view, as is usual for


R length (S.E.M.) (mm) and Range
Instar n Mouth Hooks Cephalopharyngeal Skeletona
1 133 0.021 (0.0001) (0.015-0.025) 0.085 (0.0006) (0.058-0.111)
2 193 0.039 (0.0002) (0.033-0.045) 0.149 (0.0006) (0.123-0.173)
3 234 0.063 (0.0002) (0.053-0.068) 0.233 (0.0007) (0.196-0.249)
"Cephalopharyngeal skeleton measurement included the hypopharyngeal and tentoropharyngeal sclerites.


&~~~~ &Csl $Illedll


*4* *444s4 *
4444444 *


* **



i m m I

Petitt: Distinguishing Larval Instars of Liriomyza sativae 283


A -


Fig. 2. Cephalopharyngeal skeleton including mouth hooks of first (A, B), second
(C, D), and third (E) instar larvae ofL. sativae. Partially formed mouth hooks (arrows)
posterior to the functioning mouth hooks in B and D indicate the onset of ecdysis to the
next instar. Length of mouth hooks (L-M), and length of the cephalopharyngeal skeleton
(N-O), as measured in this study, are shown in E. The same magnification was used in


1 __ ___ __..


Py- ^prw~v


jI 0. I mm

Florida Entomologist 73(2)


Ratio of Length of Cephalopharyngeal Skeleton
Ratio of
Instars L. sativae L. pictellaa L. trifoliib L. brassicaec

2/1 1.75 1.72 1.75 1.98
3/2 1.56 1.55 1.55 1.56

aOatman & Michelbacher 1958.
bParrella & Bethke 1988.
cBeri 1974.

larval Agromyzidae (Beri 1983). The relative size of each mandible could not be distin-
guished in earlier instars. The first instar mouth hooks were one-half or more as wide
as long, and only a pair of apical teeth was apparent. Occurrence of partially formed
mouth hooks along the hypopharyngeal sclerite indicated the onset of ecdysis to the
next instar (Fig. 2B and 2D).
As in other agromyzids (Beri 1973), first instar L. sativae apparently were metap-
neustic, having only the posterior pair of spiracles. The anterior spiracles of second and
third instar L. sativae are stalked with numerous spiracular bulbs.
The growth ratio of cephalopharyngeal skeletons was 1.75 between first and second
instar L. sativae and 1.56 between second and third instars. The ratios were consistent
for the species L. sativae, L. pictella, L. brassicae (Riley) and L. trifolii (Burgess), with
the exception that the ratio between first and second instars was larger in L. brassicae
(1.98) than in the other species (Table 2; Oatman & Michelbacher 1958, Beri 1974,
Parrella & Bethke 1988).
The average growth ratios between first and second and second and third instar L.
sativae mouth hooks were 1.86 and 1.62, respectively. Growth ratios between second
and third instar mouth hooks in L. andryalae Hering and L. strigata Meigen were each
1.60 (Beri 1971). The constancy of the growth ratios across species may be related to
their similar mode of feeding on leaf mesophyll tissue (Hutchinson & Tongring 1984).
Although measurements of the mouth hooks and cephalopharynqeal skeletons re-
ported in this study were made on cleared, slide-mounted larvae under a compound
microscope, 200 larvae also were classified by measurements of these structures under
a dissecting microscope. At 200x total magnification, all instars could be distinguished
by measuring either the length of the mouth hooks or cephalopharyngeal skeleton.
Because the larvae became more opaque with time in ethanol, only recently collected
larvae could be measured without clearing. Samples of 20 larvae required ca. 5 min to
classify using this method.
Rapid instar determination under the dissecting microscope makes immediate pro-
cessing of samples possible. Cohorts may be subsampled frequently, if necessary, to
determine when all individuals have reached the desired stadium for use in experimen-
tation. Availability of this method will enable studies to be conducted with particular
larval instars of L. sativae.


I acknowledge the technical assistance of Marian Coffey, Chris Halliday, and Debbie
Karan. I thank Andrew Schuerger, who assisted with photographic technique. Pauline
Lawrence and Henry Robitaille provided thoughtful criticisms of a previous draft. This
manuscript was derived from a dissertation submitted in partial fulfillment of the re-
quirements for the Ph.D. degree from the University of Florida.


June, 1990

Petitt: Distinguishing Larval Instars of Liriomyza sativae 285


BERI, S. K. 1971. Immature stages of Agromyzidae (Diptera) from India. VII.
Taxonomy and Biology of Fifteen Species of Liriomyza Mik. Oriental Ins.
(Suppl.) 1: 85-118.
BERI, S. K. 1973. Comparative morphological studies on the spiracles of larval Ag-
romyzidae (Diptera). J. Nat. Hist. 7: 481-491.
BERI, S. K. 1974. Biology of a leaf miner Liriomyza brassicae (Riley) (Diptera: Ag-
romyzidae). J. Nat. Hist. 8: 143-151.
BERI, S. K. 1983. Comparative morphological studies on the cephalopharyngeal skele-
ton of larval leaf-mining flies (Agromyzidae: Diptera). Indian J. For. 6: 301-308.
BODENSTEIN, D. 1950. The postembryonic development of Drosophila, pp. 275-367,
in M. Demerec (ed.), Biology of Drosophila. Wiley, New York.
DALY, H. V. 1985. Insect morphometrics. Ann. Rev. Entomol. 30: 415-438.
1986. Chrysocharis parksi in commercial greenhouses for the biological control
of leafminers, Liriomyza bryoniae and L. trifolii, on tomatoes; case studies and
sampling techniques. Med. Fac. Landbouww. Rijksuniv. Gent. 51(3a): 987-997.
HENDRICKSON, R. M., JR., AND S. E. BARTH. 1978. Notes on the biology of Dig-
lyphus intermedius (Hymenoptera: Eulophidae), a parasite on the alfalfa blotch
leafminer, Agromyza frontella (Diptera: Agromyzidae). Proc. Entomol. Soc.
Washington 80: 210-215.
HUTCHINSON, G. E., AND N. TONGRING. 1984. The possible adaptive significance of
the Brooks-Dyar rule. J. Theor. Biol. 106: 437-439.
IPE, I. M. 1974. Morphological, behavioral, and biological studies of Melanagromyza
obtusa (Malloch) (Diptera: Agromyzidae) on Cajanus indicus Spreng. Z. Ang.
Ent. 75: 89-98.
KAMALI, K., AND J. T. SCHULZ. 1973. Characteristics of immature stages of Gym-
nocarena diffusa (Diptera: Tephritidae). Ann. Entomol. Soc. America 66: 288-
LAWRENCE, P. O. 1979. Immature stages of the Caribbean fruit fly, Anastrepha
suspense. Florida Entomol. 62(3): 214-219.
LEIBEE, G. L. 1988. Toxicity of abamectin to Liriomyza trifolii (Burgess) (Diptera:
Agromyzidae). J. Econ. Entomol. 81: 738-740.
MASON, G. A., M. W. JOHNSON, AND B. E. TABASHNIK. 1987. Susceptibility of
Liriomyza sativae and L. trifolii (Diptera: Agromyzidae) to permethrin and fen-
valerate. J. Econ. Entomol. 80: 1262-1266.
MINKENBERG, O. P. J. M., AND J. C. VAN LENTEREN. 1986. The leafminers
Liriomyza bryoniae and L. trifolii (Diptera: Agromyzidae), their parasites and
host plants: a review. Agricultural University Wageningen Papers, 86-2.
OATMAN, E. R., AND A. E. MICHELBACHER. 1958. The melon leaf miner, Liriomyza
pictella (Thomson) (Diptera: Agromyzidae). Ann. Entomol. Soc. America 51:
PARRELLA, M. P., AND C. B. KEIL. 1984. Insect pest management: the lesson of
Liriomyza. Bull. Entomol. Soc. America 30: 22-25.
PARRELLA, M. P., V. P. JONES, AND G. D. CHRISTIE. 1987. Feasibility of parasites
for biological control of Liriomyza trifolii (Diptera: Agromyzidae) on commer-
cially-grown chrysanthemum. Environ. Entomol. 16: 832-837.
PARRELLA, M. P., AND J. A. BETHKE. 1988. Larval development and leafmining
activity of Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) Pan-Pacific En-
tomol. 64(1): 17-22.
PARRELLA, M. P., K. L. ROBB, AND P. MORISHITA. 1982. Response of Liriomyza
trifolii (Diptera: Agromyzidae) larvae to insecticides, with notes about efficacy
testing. J. Econ. Entomol. 75: 1104-1108.
PETITT, F. L. 1988. Temperature-dependent development and influence of larval in-
stars of Liriomyza sativae Blanchard on parasitization by Opius dissitus
Muesebeck. Ph.D. dissertation. Univ. of Florida. Gainesville.
ROBINSON, W. H., AND B. A. FOOTE. 1978. Biology and immature stages of Anti-
chaeta borealis (Diptera: Sciomyzidae), a predator of snail eggs. Proc. Entomol.
Soc. Washington 80(3): 388-396.

Florida Entomologist 73(2)

SPENCER, K. A. 1981. Morphological characteristics and brief taxonomic history of
Liriomyza, pp. 12-23 in D. J. Schuster (ed.), Proc. IFAS-Industry Conf. on
Biology and Control Liriomyza Leafminers II. Institute of Food and Agric. Sci-
ences. Gainesville, Florida.
TESKEY, H. J. 1981. Morphology and terminology-larvae, pp. 65-88 in J. F. McAlpine,
B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood
(eds.), Manual of Nearctic Diptera. Vol. 1. Research Branch, Agric. Canada,
Monoraph No. 27. 674 pp.
VAN ALPHEN, J. J. M., AND R. A. B. DRIJVER. 1982. Host selection by Asobara
tabida Nees (Braconidae: Alysiinae) a larval parasitoid of fruit inhabiting
Drosophila species. I. Host stage selection with Drosophila melanogaster as
host species. Netherlands J. Zool. 32: 215-231.
WESTERMAN, P. R., AND O. P. J. M. MINKENBERG. 1986. Evaluation of the effec-
tiveness of the parasitic wasps Diglyphus isaea and Chrysocharis parksi in the
experimental greenhouses for the biological control of the leafminer Liriomyza
bryoniae on tomatoes. Med. Fac. Landbouww. Rijksuniv. Gent. 51(3a): 999-


Entomology and Nematology Department
University of Florida
Gainesville, FL. USA 32611


Attraction of the pine bark beetle Hylastes salebrosus Eichhoff to the host chemicals
turpentine and ethanol, and the Dendroctonus pheromones frontalin and exo-brevico-
min, were assessed in four field trapping experiments. H. salebrosus were attracted to
turpentine alone, and the addition of ethanol, whether mixed with the turpentine or
deployed at any of three different release levels, elicited a significant increase in attrac-
tion. In another experiment incorportaing all semiochemicals tested, beetles were more
attracted to traps baited with the combination of the turpentine:ethanol mix, frontalin,
and exo-brevicomin, than to traps with the turpentine:ethanol mix only. A final pair of
experiments determined that exo-brevicomin, and not frontalin, deployed with turpen-
tine was important for eliciting attraction. H. salebrosus may produce and use exo-bre-
vicomin as an attractive pheromone, but the data also suggest that H. salebrosus can
exploit exo-brevicomin as a kairomone from other scolytid species that colonize the same
pine resource.


Atracci6n del escarabajo descortezador de pino Hylastes salebrosus Eichhoff a las
quimicas producidas por el hu6sped, aguarraz y etanol, y a las feromonas de Dendroc-
tonus frontalina y exo-brevicomina, se ensay6 en cuatro experiments de trampeo en el
campo. H. salebrosus se atrayeron a aguarraz solo, y la adici6n de etanol, ya sea mezclado


June, 1990

Phillips: Attraction of Hylastes salebrosus

con el aguarraz o puesto en combinaci6n a cualquier de 3 niveles de evaporaci6n, elicit6
un aumento significativo en la atracci6n. En otro experiment incorporando todas las
semioquimicas probadas, los escarabajos fueron mas atraidos a trampas encebadas con
la combinaci6n de mezcla aguarraz:etanol, frontalina, y exo-brevicomina, que a tram-
pas con solamente la mezcla de aguarraz:entanol. Un par final de experiments deter-
minaron que exo-brevicomina, y no frontalina, mezclado con aguarraz fu6 important
en elicitar atracci6n. H. salebrosus puede producer y usar exo-brevicomina como
feromona atrayente, pero los datos tambi6n sugieren que H. salebrosus puede explotar
exo-brevicomina como una kairomona de otras species de Scolyididae, las cuales tam-
bi6n colonizan el mismo recurso de pinos.

The bark beetle genus Hylastes Erichson is comprised of several species in North
America that breed in the lower boles and roots of diseased or moribund conifers (Wood
1982). Very little is known of the biologies and habits of these species, but their potential
as pests of conifer reproduction (Ciesla 1988) and vectors of root pathogens (Witkosky
et al. 1986) is substantial. While conducting research on the chemical ecology of Den-
droctonus terebrans (Olivier) and other pine bark beetles (e.g., Phillips et al. 1988,
1989), I observed H. salebrosus Eichhoff responding to traps baited with various
semiochemicals. Here I report results from several trapping experiments that clearly
show H. salebrosus is attracted to pine turpentine, that the attraction to turpentine is
enhanced by ethanol, and that these beetles also are attracted to a pheromone of De-
ndroctonus Erichson.

Four field experiments were conducted during 1988 in commercial slash pine, Pinus
elliottii Engelm. Var. elliottii, plantations about 20 km east of Gainesville in Alachua
Co., Florida. Traps used were 16-unit multiple funnel traps (Lindgren 1983) that were
hung from steel support stands so that the collection jars were within 15 cm of the
ground. Semiochemical release devices were hung on the outside of the third funnel
from the bottom of each trap. Experiments were set up for four to seven days as
completely randomized block designs, each with seven replicates. Treatments were
randomly assigned to traps within each block. Traps within a block were spaced 20 m
apart in a line and blocks of traps were spaced at least 40 m apart throughout the forest.
Total numbers of trapped H. salebrosus were determined, transformed via x +0.5,
subjected to analysis of variance, and means were compared using the Student-New-
man-Keuls test.
The first experiment investigated the responses of H. salebrosus to turpentine de-
ployed with different release levels of 95% ethanol. The five treatments were: 1) turpen-
tine only; 2) turpentine plus a low dose of ethanol; 3) turpentine plus a medium dose of
ethanol; 4) turpentine plus a high dose of ethanol; and 5) turpentine and ethanol released
from one container as a 1:1 mix. Ethanol doses were compared with the turpen-
tine:ethanol mix because earlier work showed that some beetle species differed substan-
tially in their responses to the mix and to undiluted turpentine with separately released
ethanol (Phillips et al. 1988). Turpentine and the turpentine:ethanol mix were evapo-
rated from 250-ml Nalgene bottles with 5 cm of a 15x1 cm cotton dental wick extending
through a 1 cm hole in the cap. Undiluted turpentine evaporated at a rate of 4.70 g/d
and the turpentine ethanol mix evaporated at 10.14 g/d; analysis of released volatiles
determined that the turpentine components (monoterpene hydrocarbons) were released
at similar levels from both bait types (see Phillips et al. 1988 for details). The turpentine
was commercially distilled from local pines and composed primarily of alpha- and beta-
pinenes (see Phillips et al. 1990). Ethanol release devices were: low ethanol, 10 ml in a
sealed 16.3x15.0 cm x 1.12 mil polyethylene sandwich bag; medium ethanol, 10 ml in an


Florida Entomologist 73(2)

open 14.8 ml glass vial (1.1 m I.D. opening); and high ethanol, 200 ml in a 250 ml
Nalgene bottle with 5 cm of a 15x1 cm cotton dental wick extending through the cap.
Evaporation rates of ethanol from these devices were: low, 3.18 mg/hr; medium, 25.84
mg/hr; and high, 596.40 mg/hr.
The second experiment, designed initially to study the responses of D. terebrans
(Phillips unpublished), examined the responses of H. salebrosus to the turpen-
tine:ethanol mix and the bark beetle pheromones frontalin and exo-brevicomin. The five
treatments were: 1) turpentine:ethanol only; 2) turpentine:ethanol plus frontalin re-
leased from three glass capillaries; 3) turpentine:ethanol plus three capillaries of fronta-
lin and one capillary of exo-brevicomin; 4) turpentine:ethanol plus three capillaries of
frontalin and three capillaries of exo-brevicomin; and 5) turpentine:ethanol plus three
capillaries of frontalin and five capillaries of exo-brevicomin. Source, purities, release
devices, and release rates for the synthetic, racemic pheromones are given in Phillips
et al. 1990.
Because the second experiment found that the Dendroctonus pheromones were at-
tractive to H. salebrosus, the third and fourth experiments were designed to determine
the roles of frontalin and exo-brevicomin separately. Both experiments utilized a low
release rate of undiluted turpentine so that any attractive effects of the synthetic
pheromones would not be overwhelmed by the attraction to host odors. Turpentine was
evaporated from an open 14.8 ml glass vial (1.1 cm I.D. opening) at a rate of about 180
mg/d, as determined in the laboratory at 30 C. The third experiment assessed the
responses of H. salebrosus to traps baited with 1) turpentine only, and 2) turpentine
plus three capillaries of frontalin. The fourth experiment compared responses to 1)
turpentine only, and 2) turpentine plus three capillaries of exo-brevicomin.


Turpentine alone was attractive to Hylastes salebrosus, and the addition of ethanol
at any release level markedly increased this attraction (Fig. 1). Earlier studies failed
to catch any H. salebrosus or other phloem-feeding bark beetles at traps baited with
ethanol only (Phillips et al. 1988) or at unbaited traps (unpublished). Therefore, ethanol
acts synergistically to increase the attraction of H. salebrosus to turpentine. This syn-
bergistic effect of ethanol is similar to that found for D. terebrans (Phillips et al. 1988),
Hylobius pales Herbst (Curculionidae) (Fatzinger 1985), H. abietis (L.) (Tilles et al.
1986), Monochamus titillator (F.) (Cerambycidae) (Fatzinger et al. 1987), and other
pine-infesting beetles (e.g., Klimetzek et al. 1986, Ch6nier & Philogene 1989). Varied
doses of ethanol affected responses of other beetles to host odors and pheromones in
other studies (e.g., Klimetzek et al. 1986), but different doses of ethanol had no signif-
icant effect on the responses of H. salebrosus in this study. I believe this is the first
clear demonstration of a turpentine-ethanol synergism for a species of Hylastes. Wit-
kosky et al. (1987) reported that H. nigrinus (Mannerheim) was more attracted to
alpha-pinene than to ethanol, but they did not combine the two to test for synergism
or enhancement. Chenier & Philogene (1989) caught very low numbers of H. porculus
Erichson and did not detect a significant difference between responses to monoterpenes
and ethanol, either singly or in combination.
Turpentine, which is comprised primarily of monoterpene hydrocarbons, is the vol-
atile fraction of pine oleoresin and is representative of odors emanating from cut or
injured pines. Ethanol is produced by stressed, injured, or decomposing conifers and
other woody plants (Moeck 1970, Kimmerer & Kozlowski 1982) and is attractive alone
to many scolytids and cerambycids (e.g., Roling & Kearby 1975, Montgomery & Wargo
1983). Turpentine, therefore, would be a general cue from a potential host for pine-
breeding insects, but ethanol with host terpenes would present a more defined signal


June, 1990

Phillips: Attraction of Hylastes salebrosus



T+LOW ETOH/////////// B

T+MED ETOH ////////////// B



0 10 20 30

Fig. 1. Mean responses and standard errors of Hylastes salebrosus in the first
experiment; TURP and T=Turpentine, ETOH=ethanol deployed at various doses
(low, medium, high), T:E = a 1:1 mix of turpentine and ethanol. Means followed by the
same letter are not significantly different (P>0.05, Student-Newman-Keuls test;
F=6.20, df= 10, 24, P=0.001).

regarding the physiological stress or state of decompostition of the host. This scenario
of host selection, in which terpenes and ethanol signify a host of a particular condition,
applies to H. salebrosus, which colonizes dead pine material or cut stumps and roots of
recently killed trees (Wood 1982).
Traps baited with the combination of turpentine:ethanol, frontalin, and a high dose
of exo-brevicomin in the second experiment caught significantly more H. salebrosus
than traps baited with turpentine:ethanol only (Fig. 2). In the third experiment beetles
did not respond differently to traps baited with turpentine or turpentine plus frontalin,
but the fourth experiment demonstrated that turpentine plus exo-brevicomin was signif-
icantly more attractive than turpentine only (Fig. 3). The low number of beetles caught
in the third and fourth experiments can be attributed to the relative lower level of
turpentine released (and perhaps the lack of ethanol) compared with other experiments.
Data from the third and fourth experiments suggest that responses of H. salebrosus in
the second experiment (Fig. 2) were not strongly affected by frontalin, but that exo-bre-
vicomin was primarily responsible for the significant attraction to the combination of
turpentine:ethanol, frontalin, and the high dose of exo-brevicomin. Previous studies
(Phillips et al. 1989, unpublished) failed to attract any H. salebrosus to traps baited
with exo-brevicomin only, therefore I assume that the attraction observed here to tur-
pentine with exo-brevicomin, like similar phenomena in other bark beetles (Borden
1982), resulted from a synergism of the two materials deployed together.
There are at least two explanations for why H. salebrosus was attract to exo-bre-
vicomin in my experiments. First, one or both sexes of H. salebrosus may produce and
utilize exo-brevicomin as an attractant pheromone. Exo-brevicomin is a common
pheromone in several species of the closely related genus of Dendroctonus, and also
occurs in other scolytids (Borden 1982). Support of this hypothesis would require, among
other data, experimental proof of intraspecific pheromone-based attraction in H. saleb-
rosus, and subsequent chemical identification of exo-brevicomin from the pheromone-

Florida Entomologist 73(2)






0 5 10 15 20


Fig. 2. Mean responses and standard errors of Hylastes salebrosus in the second
experiment; T:E=a 1:1 mix of turpentine and ethanol, FRONT and F=frontalin,
EXO = exo-brevicomin deployed at various doses (lx, 3x, 5x). Means followed by the
same letter are not significantly different (P>0.05, Student-Newman-Keuls test;
F=2.81, df= 10, 24, P=0.018).

producing beetles. A second, and more accessible, explanation for the attraction elicited
by exo-brevicomin is that this compound represents a kairomone produced by co-attack-
ing species of bark beetles. H. salebrosus may exploit the exo-brevicomin produced by
other species, such as D. terebrans (Phillips et al. 1989), which would have made the
initial effort and risk of locating and attacking a tree. Exo-brevicomin, therefore, would
signify to H. salebrosus a suitable host for colonization (i.e., one dying or being killed
by other bark beetles) and also a place for locating potential mates. Interspecific exploi-
tation of chemical signals is common in bark beetles and their associates (Borden 1982).

5 5
S 4 -4

3 3

C 2. 2
6 1 1

W 0 0

Fig. 3. Mean responses and standard errors of Hylastes salebrosus in the third (A)
and fourth (B) experiments; LOW T=a low dose of undiluted turpentine,
FRONT = frontalin, EXO = exo-brevicomin. Differences between means for each exper-
iment were assessed by analysis of variance (NS, P=0.776, F=0.084, df=l, 12; **,
P=0.032, F =5.815, df= 1, 12).


June, 1990

Phillips: Attraction of Hylastes salebrosus

It is possible, of course, that H. salebrosus both produces exo-brevicomin as a
pheromone and also responds to it opportunistically as a kairomone. Such a pheromone-
kairomone system has been suggested for D. terebrans and D. frontalis Zimmermann,
both of which commonly co-attack the same host trees and produce and respond to
frontalin (Payne et al. 1987).


I am grateful to Tom Atkinson for initially identifying H. salebrosus from my sam-
ples, for writing the Spanish translation of the abstract, and for his review of the
manuscript. I also thank John Foltz and Bob Wilkinson for their reviews of the manu-
script. This research was supported by a grant from the USDA Competitive Research
Grant Program, 87-CRCR-1-2491. This paper is Florida Agricultural Experiment Sta-
tion Journal Series no. R-00282.


BORDEN, J. H. 1982. Aggregation pheromones. pp. 74-139 in J. B. Mitton, and K. B.
Sturgeon (eds.), Bark beetles in North American conifers. University of Texas
Press, Austin. 527 pp.
CHENIER, J. V. R., AND B. J. R. PHILOGENE. 1989. Field responses of certain forest
Coleoptera to conifer monoterpenes and ethanol. J. Chem. Ecol. 15: 1729-1745.
CIESLA, W. M. 1988. Pine bark beetles: a new pest management challenge for Chilean
foresters. J. Forestry. 86: 27-31.
FATZINGER, C. W. 1985. Attraction of the black turpentine beetle (Coleoptera:
Scolytidae) and other forest Coleoptera to turpentine-baited traps. Environ. Ent.
14: 768-775.
Trans-verbenol, turpentine, and ethanol as trap baits for the black turpentine
beetle, Dendroctonus terebrans, and other forest Coleoptera in North Florida.
J. Entomol. Sci. 22: 201-209.
KIMMERER, T. W., AND T. T. KOZLOWSKI. 1982. Ethylene, ethane, acetaldehyde,
and ethanol production by plants under stress. Plant Physiol. 69: 840-847.
KLIMETZEK, D., J. KOHLER, J. P. VITE, AND U. KOHNLE. 1986. Dosage response
to ethanol mediates host selection by "secondary" bark beetles. Naturwis-
senschaften 73: 270-272.
LINDGREN, B. S. 1983. A multiple funnel trap for scolytid beetles (Coleoptera). Cana-
dian Entomol. 115: 299-302.
MOECK, H. A. 1970. Ethanol as a primary attractant for the ambrosia beetle
Trypodendron lineatum (Coleoptera: Scolytidae). Canadian Entomol. 192: 985-
MONTGOMERY, M. E., AND P. M. WARGO. 1983. Ethanol and other host derived
volatiles as attractants to beetles that bore in hardwoods. J. Chem. Ecol. 9:
W. FRANKE, AND J. P. VITE. 1987. Kairomonal-pheromonal system in the
black turpentine beetle, Dendroctonus terebrans (01.). J. Appl. Entomol. 103:
SON, AND J. L. FOLTZ. 1988. Synergism of turpentine and ethanol as attrac-
tants for certain pine-infesting beetles (Coleoptera). Environ. Entomol. 17: 456-
ary attraction and field activity of beetle-produced volatiles in Dendroctonus
terebrans. J. Chem. Ecol. 15: 1513-1533.

292 Florida Entomologist 73(2) June, 1990

A. C. OEHLSCHLAGER. 1990. Response specificity of Dendroctonus terebrans
(Coleoptera: Scolytidae) to enantiomers of its sex pheromones. Ann. Entomol.
Soc. Am. 83: 251-257.
ROLING, M. P., AND W. H. KEARBY. 1975. Seasonal flight and vertical distribution
of Scolytidae attracted to ethanol. Canadian Entomol. 107: 1315-1320.
gism between ethanol and conifer host volatiles as attractants for the pine weevil
Hylobius abietis (L.) (Coleoptera: Curculionidae). J. Econ. Entomol. 79: 970-974.
WITKOSKY, J. J., T. D. SCHOWALTER, AND E. M. HANSEN. 1986. Hylastes nigrinus
(Coleoptera: Scolytidae), Pissodes fasciatus and Steremnius carinatus (Col-
oeoptera: Curculionidae) as vectors of black-stain root disease of Douglas-fir.
Environ. Entomol. 15: 1090-1095.
WITKOSKY, J. J., T. D. SCHOWALTER, AND E. M. HANSEN. 1987. Host-derived
attractants for the beetles Hylastes nigrinus (Coleoptera: Scolytidae) and
Steremnius carinatus (Coleoptera: Curculionidae). Environ. Entomol. 16: 1310-
WOOD, S. L. 1982. The bark and ambrosia beetles of North and Central America
(Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Natur. Mem. 6,
Brigham Young University, Provo, Utah.


U. S. Vegetable Laboratory
2875 Savannah Highway
Charleston, S.C. 29414

Insect Attractants, Behavior, and Basic Biology Research Laboratory
P.O. Box 14565
Gainesville, FL 32604


Capture of male banded cucumber beetles Diabrotica balteata LaConte in sex
pheromone-baited traps or in virgin female baited traps peaked at the beginning of the
scotophase (2100-2200h) and declined thereafter. Virgin females (1 or 2) were not as
effective as the racemic synthetic sex pheromone (0.5mg/septum) in attracting males.
the most efficient design of capturing males was the Sentry wing trap and no differences
in captures were found among traps placed 25.4, 50.8 or 101.6cm from the ground. Field
tests showed that a rubber septum bait containing 0.5mg of the pheromone was attrac-
tive to males for a 12 month period and high captures occurred in August and Sep-
tember. Males were collected in pheromone traps from October through February in
Charleston, SC.

Schalk et al.: Diabrotica balteata Sex Pheromone



La capture de machos del escarabajo Diabrotica balteata LeConte en trampas
cebadas con feromonas sexuales o en trampas cebadas con hembras virgenes, tuvo el
auge al comienzo de la escotofase (2100-2200h) y despuds decline. Hembras virgenes (1
6 2) no fueron tan efectivas en atraer machos como la feromona sexual sintetica rac6mica
(0.5mg/septum). El disefio mAs efectivo para capturar machos fuh la trampa de ala
Sentry, y no hubo diferencia en la capture entire trampas puestas a 25.4, 50.8, o a 101.6
cm del suelo. Pruebas de campo demostraron que la trampa cebada con septum de goma
conteniendo 0.5 mg de la feromona, fue atractiva para los machos por un period de 12
meses, ocurriendo muchas captures en Agosto y Septiembre. Se colectaron machos en
trampas de feromona de Octubre a Febrero en Charleston, SC.

The banded cucumber beetle (Diabrotica balteata LeConte) is a polyphagous species
which is a serious pest of root crops in the southern states (Metcalf et al. 1962, Krysan
& Branson 1983). Control of the soil inhabiting larvae of this beetle is difficult because
of the nonpersistent nature of the currently registered insecticides. However, it may
be possible to reduce larval populations by controlling adult beetles, as they are easily
killed with foliar insecticides. The recent identification and synthesis of the banded
cucumber beetle sex pheromone (6,12-dimethylpentadecan-2-one) (Chuman et al. 1987)
could enhance adult control programs by making possible more effective timing of insec-
ticide treatments. The pheromone may also be useful for direct control of adults as a
component of an IPM program. This paper reports basic information on the diel and
seasonal responses of males to sex pheromone, the longevity of a pheromonal formula-
tion, and the efficacy of several types of traps.


The racemic pheromone used in the tests of 1987 was synthesized according to
Chuman et al. (1987). The synthetic pheromone used in the 1988 tests was racemic
6,12-dimethylpentadecan-2-one which was purchased from Fuji Flavor Co., Ltd.,
Tokyo, Japan. Its structure was confirmed by mass spectroscopy (Chuman et al. 1987)
and its purity assessed by gas chromatography on two 50 m x 0.25 mm fused silica
capillary columns, one coated with OV101 and the other with CPS-2 (Quadrex Scientific,
New Haven, CT). The pheromone comprised 84% of the synthetuc material. The re-
mainder consisted of one unknown impurity of 12% and several minor (1% or less)
impurities. Rubber septa (A. H. Thomas #8753-D22) were extracted for 48 h with
methlylene chloride in a soxhlet extractor to remove any impurities and dried at ambient
temperature in a fume hood. The synthetic pheromone, in the amounts indicated for the
various experiments, was loaded into the septa by pipetting 100 ul of hexane solution
into the well of the large end of each septum. Septa were allowed to equilibrate in a
fume hood at ambient temperature for 48 h after loading.
The tests were conducted at the U. S. Vegetable Laboratory, Charleston, SC.
The first experiment was conducted in 1987 to determine the diel periodicity of male
response to synthetic pheromone-baited traps and to traps baited with virgin female
beetles. Traps for these tests were 450 ml white styrofoam cups externally covered with
insect trapping adhesive and suspended upside down on wooden stakes with cup orifices
15 cm above the plant canopy. Traps were baited with rubber septa containing 0.5 mg
of racemic pheromone or with nylon mesh cages (5Lx 2D cm, containing a moist dental
role wick) confining 1 or 2, 10 to 26 day-old virgin females. Cuthbert & Reid (1964)
found that the mean attractiveness to males for recently emerged females was 10 days,
however, the range of attractiveness was 4 to 79 days and depended on the females.

Florida Entomologist 73(2)

These baits were placed on top of the inverted cup. Five of each type of trap were
placed 15 m apart consecutively in the center of sweet potato fields. Treatments were
placed in the fields after 1500 h and data recorded every 30 min from 1600 to 2400 and
from 0800 to 1500 h the following day. Four tests were conducted in July (July 20, 21,
29 and 30), and 3 in September (September 13, 15, and 16) which assessed male response
to both the pheromone and virgin females. In another study male response to only the
pheromone was investigated on July 29, 30, August 3, 4 and September 1, 2. The onset
of the scotophase occurred about 2130 h on 20 July and 2100 h on 1 September. Average
temperature for the test period ranged from 25 to 31C with only traces of moisture
reported for 1 September. Average wind speed did not exceed 22 k/h. Captured adults
were counted, removed from the trap and brought into the laboratory for sex confirma-
In a second test, conducted in 1988, six trap designs were compared with the 450-ml
cup trap for efficacy in capturing males: Sentry wing (Pherocon IC, Albany Interna-
tional Co.); a boll weevil trap (Techical Precision Plastics); Jansson sweetpotato weevil
trap (Jansson, et al. 1989); Multi-pher trap (Bio-control Services); Japanese beetle-Bio-
lure trap (Reuter Laboratories, Inc.); and Delta-Biolure trap (Consep Membranes,
Inc.). Each trap was baited with a septum containing 300 ug of pheromone and an
unbaited 450 ml cup trap was included. Three trapping lines, each with 8 trap locations,
were established in a field of sweet corn at the silk stage on 22 June (corn 1.5 m high,
with traps placed 61 cm from the ground). The trap locations within each line were 36
m apart and the trap lines were 45 m apart. Each trap design was randomly assigned
to a trap location within each line to establish a randomized complete-block experiment.
The captured males were counted and removed from the traps daily and each trap was
then moved to the next location in its trap line for 8 days. The trap captures were
transformed to X+ 0.1 to stabilize the variance and analyzed using PROC ANOVA
(SAS 1985) to determine effects of trap design, block and position.
Mean separation for trap design was by the SNK (Student-Newman-Keuls) proce-
dure. During the test, the average daily temperature was 27C with trace amounts of
precipitation on June 24 and 27. Wind speed averaged less than 22 k/h during this
period. The third test, conducted on 1 August, 1988, determined the effect of placing
traps at various heights in a snap bean (45 cm tall) field (spacing was 15 m between
traps and 8 m between replicates) by positioning the wing design at 25.4, 50.8 and 101.6
cm above the ground. Each trap contained 300 ug of pheromone. Trap data were col-
lected daily for 12 consecutive days and trapped insects were removed daily. The aver-
age daily temperature and precipitation for the test period was 28.4C and 1.8 cm
respectively. Wind speed did not exceed 24 k/h. The experimental design was a 3 x 3
Latin square.
In the final test, pheromone activity was investigated by leaving five baited traps
(450 ml cups, 0.5 mg pheromone/trap) with the original septa continuously in the field,
spaced 15 m apart and positioned on top of ditch banks (91 cm above ground), from July
1987 through May 1988. Data were recorded 3 times/week and traps were periodically
cleaned of beetles or replaced.


Peak captures of males in pheromone baited traps and virgin female traps occurred
during the scotophase between 2100 through 2400h and subsequently declined (Fig. 1).
Howard (1982) found similar activity with sweep net captures of unsexed adult banded
cucumber beetles. Cuthbert (1964) stated that females released their pheromone during
at least a portion of the night. Chuman et al. (1987) captured large numbers of males
during both morning and afternoon tests in south Florida. They attributed this pattern


June, 1990

Schalk et al.: Diabrotica balteata Sex Pheromone 295



o 6000
S--- Males Captured

a 4000



July Aug. Sept. Oct. Nov. Dec. Jan. Feb. March April May June
1987 1988

Fig. 1. Field evaluation of the pheromone racemicc 6,12-dimethylpentadean-2-one)
(0.5 mg/septa/trap) used continually for a 12 month period to capture males of Diabrotica
balteata LeConte, 1987-1988.

of response to cool night temperatures in November which presumably inhibited beetle
flight. Subsequent trapping tests during the summer months in Florida (McLaughlin,
unpublished) revealed a male response similar to that presented here for South Carolina.
These findings suggest that males adjust their response under the influence of environ-
mental cues, however, we do not know if female pheromone release is similarly affected.
Virgin females were not as effective as the synthetic pheromone as demonstrated by
total numbers of captured males per season (Fig. 1, Table 1).
The most effective trap design for capturing males was the wing type (Table 2). The
Delta, Multi-pher trap and 450 ml cup designs were intermediate in male capture while
the least effective were the Japanese beetle, Jansson and boll weevil traps. The unbaited
450 ml cup without the pheromone, did not trap any males.


Males Captured
Treatment (Mean SE/day)
(date) Pheromone Female

9/13 94.8 13.3 38.8 7.6
9/15 64.6 4.3 59.4 14.3
9/16 84.4 10.2 24.6 8.2
Total 243.8a2 122.8b

'Traps baited after 1530h each day; 5 traps per treatment; females were 10-26 days old.
'Means between columns (total captures) not followed by the same letter are significantly different at the 1% level
(t-test for paired replicates).

Florida Entomologist 73(2)

June, 1990


Males captured
Trap design (Mean + SE/day)'

Wing 11.3 2.7a
Delta 7.6 2.8b
Multitrap 5.0 + 1.4b
450ml cup 4.9 1.4b
Japanese beetle 2.1 0.6c
Jansson 2.1 0.5c
Boll weevil 1.5 0.4c
450ml cup (unbaited) 0.0 + 0.0d

'Means not followed by a common letter are significantly different. Student-Newman-Keuls procedure, alpha =
0.05, F = 26.7; df = 7.32; P>F = 0.0001.

There were no significant differences in males captured with wing traps positioned
at heights of 25.4, 50.8 or 101.6 cm. (means of 8.6 SE 12.1, 10.5 SE 11.8 and 9.6
SE 10.6 males, respectively, per day). This trapping was in a field of snap bean with
canopy height of about 45 cm. These findings indicate that in low growing crops the
traps can be elevated above the canopy so they may be more easily located by field
The 0.5 mg baits attracted males continuously from July 1987 through May 1988.
Peak captures were observed during the months of August and September of 1987 with
a decline into the cooler seasons (Fig. 2). This agrees with the seasonal densities that
Elsey (1988) found on cucumber and zucchini squash. However, the effectiveness of the
bait was probably reduced due to the length of use. The important observation is capture
of males during the months of January, February, and March which demonstrates that

I pheromone
o Virgin females Testing per

lod from 7/29-9/2

Testing Period from 7/20-7/31

m L .- l_. -
16-17 17-18 18-19 19-20 20-21 21-22 22-24 24-8 9-10 10-11 11-12 12-13 13-14 14-15
a 0

Day 1

Day 2

Hour Collected (DST)

Fig. 2. Numbers of male Diabrotica balteata captured over a 24 h (1600 h through
1500 h) period in traps baited with 0.5 mg of racemic 6,12-dimethylpentadecan-2-one or
virgin females.





Schalk et al.: Diabrotica balteata Sex Pheromone

adults can survive the winter in Charleston, SC. Elsey (1988) reported similar findings
and also suggested that Charleston, SC. may be one of the northernmost enclaves for
the banded cucumber beetle.
The larval stage of this insect is very difficult to control because of the loss of
persistent soil insecticides, however, the adults can be effectively controlled with an
economical chemical such as Sevin. The male insect's sensitivity to the pheromone
suggests that it may be useful for integrated pest management. It can be employed as
bait in wing traps to detect early infestations, to monitor established populations and
to assist correct timing of insecticide applications relative to population densities of
economic importance. Since the insect is a pest of seasonal crops the pheromone will
probably have little value in mass trapping because of the lack of crop continuity. Its
value for mating disruption seems poor because of long adult life which enhances greater
likelihood of copulation with females.


M. SCHALK, AND J. H. TUMLINSON. 1987. Identification of female-produced
sex pheromone, from banded cucumber beetle, Diabrotica balteata LeConte (Col-
eoptera:Chrysomelidae). J. Chem. Ecol. 13: 1601-1616.
CUTHBERT, F. P., JR., AND W. J. REID, JR. 1964. Studies of sex attractant of
banded cucumber beetle. J. Econ. Entomol. 57: 249-250.
ELSEY, K. D. 1988. Cucumber beetle seasonality in coastal South Carolina. Envir.
Entomol. 17: 496-502.
HOWARD, F. W. 1982. Diurnal rhythm in Cylas formicarius elegantulus and some
other arthropods in a sweet potato field. Florida Entomol. 65: 194-195.
JANSSON, R. K., R. R. HEATH, AND J. A. COFFELT. 1989. Temporal and spatial
patterns of sweet potato weevil (Coleoptera:Curculionidae) counts in pheromone-
baited traps in white-fleshed sweet potato fields in southern Florida. Envir.
Entomol. 72: (in press).
KRYSAN, J. L., AND T. F. BRANSON. 1983. Biology, ecology and distribution of
Diabrotica. Proc. Inter. Maize virus disease colloquium and workshop. The Ohio
Univ., Ohio Agricultural Research and Development Center, Wooster. 1-7.
METCALF, C. L., W. P. FLINT, AND R. L. METCALF. 1962. Destructive and useful
insects. McGraw Hill Book Co. Inc. N. Y., San Francisco, Toronto, London.
SCHALK, J. M. 1986. Rearing and handling of Diabrotica balteata, pp. 49-56 in J. L.
Krysan and T. A. Miller [eds.], Methods for the study of pest Diabrotica.
Springer-Verlag. New York, Berlin, Heidelberg, Tokyo. 260 p.


Florida Entomologist 73(2)


Research Department
United States Sugar Corporation
Clewiston, Florida 33440


Stand reductions in plant cane caused by Melanotus communis (Gyll.) were quan-
tified in small-plot tests with 2 commercial clones and in a large-plot study with 1
commercial clone. A standard measure of the number of wireworms per 1.5 row-meters
was used. Infestation levels studied ranged from 0 up to 12 M. communis per 1.5
row-m. Stand reductions per wireworm per 1.5 row-meters ranged from 6.2 to 7.8% at
12 weeks after planting based on regression analyses. Tillering during the growing
season compensated to some degree for early stand losses but, at harvest, stalks from
plots infested by wireworms tended to weigh less. Yield in terms of the total weight of
cane in plots at harvest was reduced by wireworms. In the large-plot study, the weight
of cane harvested per hectare was reduced by 5.9 metric tons or by 3.8% per wireworm
per 1.5 row-meters (r = .92). Wireworms did not cause reductions in sucrose levels in
juice. However, because cane weight at harvest was reduced by 3.8% per wireworm
per 1.5 row-meters, the amount of sugar produced was reduced by the same amount.
Stand reductions in plant cane caused by 8 to 12 wireworms/1.5 row-meters carried over
into the first ratoon crop, which resulted in lower yield of the ratoon crop.


Se cuantificaron las p6rdidas en la reducci6n de plants en caria de plantilla causadas
por Melanotus communis (Gyll.) en el campo en pequefias parcelas con dos clones
comerciales y en un studio con una parcela grande de un clon commercial. Se us6 la
media patr6n de gusanos de alambre en 1.5 metros de zurco. La disminuci6n en el
nimero de plants por cada gusano de alambre en 1.5 metros de zurco fluctu6 entire 6.2
y 7.8% basado sobre datos de un analisis de regresi6n del nimero de plants contadas
a las 12 semanas despu6s de la siembra. El macollamiento durante la temporada de
crecimiento compens6, hasta cierto punto, la p6rdida de plants, pero los tallos de
parcelas infestadas por gusanos de alambre tendieron a pesar menos cuando se cosecha-
ron. El rendimiento en terminos del peso total de la cana en parcelas fue reducido por
los gusanos de alambre. En el studio de la parcela grande, el peso por hectarea de la
cafia cosechada se redujo un 3.8% por cada gusano de alambre en 1.5 metros de zurco
(r = .92). Los gusanos de alambre no causaron reducci6n en los niveles de sacarosa del
jugo en estas pruebas. Sin embargo, debido a que el peso de la cafa fue reducido un
3.8% por cada gusano de alambre en 1.5 metros de zurco, la cantidad de azficar se redujo
por la misma cantidad. La reducci6n de plants en caila plantilla causada por 8 a 12
gusanos de alambre en 1.5 metros de zurco en el studio de la parcela grande, tambi6n
afecto el cultivo de primer retofio.

The wireworm Melanotus communis (Gyll.) is an important pest of sugarcane grown
in southern Florida (Gifford 1964, Hall 1988). M. communis is a more common pest of
cane grown in soils with a moderate to high content of organic matter (Cherry & Hall


June, 1990

Hall: Losses to Wireworms in Sugarcane

1986). Feeding damage by this wireworm to seedpieces and to young developing shoots
results in stand losses (Ingram et al. 1938, 1951). The establishment of a good stand is
critical in sugarcane production because stand is directly related to yield as well as to
successful ratooning. M. communis is primarily important as a pest of sugarcane during
the plant-cane crop; one study indicated that significant stand losses can occur at a level
of about three wireworms per 1.5 row-meters in plant cane during the first six weeks
of stand development (Hall 1985). Wireworms infesting ratoon cane have generally had
less of an impact on stand, but significant losses caused by M. communis in ratoon cane
may occasionally occur (personal observations).
The purpose of research presented in this publication is to further elucidate the
importance of M. communis as a pest of plant cane in Florida. Different population
levels of the wireworm were introduced into plant-cane plots, and the effects of
wireworms on stand and yield were investigated. After plant cane was harvested, a
first-ratoon crop was grown in order to determine if wireworm damage during plant
cane had any effect on stand and yield of the following crop.


Stand reductions caused by M. communis infestations in young plant cane were
assessed in two experiments conducted on muck (>50% organic matter) soil near Clewis-
ton in Palm Beach County. Wireworms were collected from commercial sugarcane fields
in late December and January and introduced at planting time into plots of sugarcane
at various infestation levels per 1.5 row-meters. This was a convenient basis for
wireworm densities and could easily be mathematically converted to population levels
per field or per hectare. Several days or more were required to collect enough
wireworms for each test. Therefore, some wireworms had to be maintained in large
pans containing soil and wheat seeds in a laboratory until enough had been collected.
Because there was variability in body size and age of wireworms collected from commer-
cial cane fields, a mix of these wireworms was introduced into each test plot. When
wireworms were introduced into test plots, they were spaced evenly apart along the
cane rows; wireworm populations generally appear to be somewhat randomly or un-
iformly dispersed in cane fields at planting time, at least within localized areas compar-
able in size to the plots used in these studies (personal observations). In both experi-
ments, test plots were planted at a rate of 20 eyes per 1.5 row-meters in soil that had
been fallow for nine months, and an unplanted 4.5 meter buffer zone was left around
each plot. Bait traps similar to those used by Ward & Keaster (1977) indicated no M.
communis were present in soil at the test sites three weeks before planting.
In a 1987 small-plot test, wireworms were introduced at 0, 2, 4, 6, or 8 per 1.5
row-meters into single-row, 1.5-m plots immediately after planting on January 21. An
appropriate number of evenly-spaced, small holes that extended down to seed piece
level were made using a soil probe along each row; a wireworm was dropped into each
hole and covered. One set of plots was planted with the commercial clone CL 61-620
and one was planted with the commercial clone CL 59-1052. A randomized block design
with ten replications of each infestation level was used for each clone. Stand counts in
each plot were made at 9, 12, and 36 weeks after planting. At harvest during the first
week of November, a sample of ten stalks from each plot was taken to a laboratory,
stripped of leaves, topped above the uppermost hardened internode, and weighed. Each
10-stalk sample was then passed through a research 3-roll mill to extract and collect
juice. The juice was analyzed to determine the yield of 96 sucrose as a percentage of
cane weight. Factors developed by Bourne (1968) were used in the calculations. The
remaining stalks in each plot were burned, hand cut, topped and weighed.


Florida Entomologist 73(2)

June, 1990

In a 1988-90 large-plot study, 0, 4, 8, or 12 M. communis wireworms per 1.5 row-me-
ters were introduced just after planting on January 14 into plots of CL 61-620 that were
2 rows by 13.1 m (1.5 meter row-spacing). A randomized complete block design with
four replications was used. A shallow trench (about 5 cm deep) was made above the
seedpieces along each row, wireworms were placed into the trench such that they were
uniformly spaced along each row, and then the wireworms were covered with soil.
Stand losses were assessed at 8 and 12 weeks after planting. The yield of 96 sucrose
as a percentage of cane weight was determined in each plot on January 20, 1989, using
core-punch samples of juice (Bourne 1968). The plots were burned to remove leaves and
tops, hand cut, and weighed during the first week of March, 1989. The plots were
allowed to ratoon naturally to determine if stand and yield losses caused by wireworms
during the plant cane crop carried over to the ratoon crop. Stand of first-ratoon cane
in each plot was assessed on August 17, 1989. The first-ratoon cane was burned, hand
cut, and weighed during the third week of February, 1990.
Statistical analyses on data from these studies were conducted using PROC ANOVA
along with REGWQ (P = .05) and PROC GLM (SAS Institute 1985).


Analyses of variance of data from small plots of CL 61-620 and CL 59-1052 indicated
that an infestation level of six M. communis wireworms per 1.5 row-m in either variety
at planting time significantly reduced stand during the first 12 weeks of growth (Table
1). Stand development after 12 weeks compensated to some degree for early stand
losses caused at six wireworms per 1.5 row-meters. The stand difference between CL


R Stand per plot At harvest
after planting R weight (kg)
per 1.5 Per Cane
Clone row-meters 9 Wks 12 Wks 36 Wks stalk per plot

CL61-620 0 5.0a 6.4a 32.3a 1.39a 44.7a
2 6.0a 6.6a 33.2a 1.41a 46.4a
4 5.2a 5.8ab 31.3ab 1.36a 42.7a
6 4.3ab 4.4bc 30.2ab 1.34a 40.4a
8 2.5c 3.1c 24.3b 1.22a 29.8b
F 4.7 6.1 2.9 2.2 5.8
P>F .0037 .0007 .0361 .0876 .0011
CL59-1052 0 4.4a 4.8a 19.7a 1.31a 26.3a
2 3.3ab 3.9ab 16.la 1.35a 22.0a
4 3.2ab 3.5ab 16.6a 1.35a 22.6a
6 2.2bc 2.8bc 16.0a 1.24a 19.9a
8 1.4c 1.6c 7.4b 1.26a 9.4b
F 7.0 5.7 6.9 1.0 6.2
P>F .0003 .0012 .0003 .4029 .0007

'For each clone, means in the same column followed by the same letter are not significantly different using the
Ryan-Einot-Gabrie-Welsch multiple range test. The error d.f. for each ANOVA was 36 except for analyses on
individual stalk weights, which had 35 d.f. for CL 61-620 and 30 for CL 59-1052. These d.f. were ower because eight
wireworms per 1.5 row-m sometimes completely eliminated stand.


Hall: Losses to Wireworms in Sugarcane 301

61-620 and CL 59-1052 after 36 weeks in plots not infested was normal. Regression
analyses of data collected 12 weeks after planting indicated stand was reduced by 6.8%
per wireworm per 1.5 row-m (r = .57, F = 23.3, P > F = .0001) in plots of CL 61-620
and by 7.8% per wireworm per 1.5 row-m (r = .58, F = 22.8, P > F = .0001) in plots
of CL 59-1052. Wireworms tended to cause larger stand reductions in CL 59-1052,
probably because this clone germinates slower and produces fewer stalks per ha. No
significant differences were found among wireworm levels in the average weight of
individual stalks of each clone at harvest, but stalk weights tended to be lower in plots
infested by six or eight wireworms. Based on ANOVA, reductions in kilograms of cane
produced per plot were significant at eight wireworms per 1.5 row-meters in each clone.
Regression analyses indicated the weight of cane harvested per plot was reduced by
1.8 kg per wireworm per 1.5 row-m in CL 61-620 (r = .48, F = 14.7, P > F = .0004)
and by the same amount in CL 59-1052 (r = .48, F = 14.6, P > F = .0004). Although
yield reductions caused by wireworms were similar in the two clones, the percentage
loss in CL 59-1052 was greater due to its lower yield potential in this test. No differences
among wireworm infestation levels were found in either variety with respect to sucrose
levels in juice.
In the large-plot test with CL 61-620, stand reductions caused by M. communis
were evident at eight weeks after planting (Table 2). Significant reductions occurred at
four wireworms per 1.5 row-meters. Increasing the wireworm density from four to
eight per 1.5 row-meters significantly increased stand losses based on ANOVA of data
from 8 and from 12 weeks after planting. Regression analyses of data collected 12 weeks
after planting indicated stand was reduced by 6.2% per wireworm per 1.5 row-m (r =
.97, F = 214.2, P > F = .0001). Yield in terms of tonnage of cane per hectare was
significantly reduced by as few as four wireworms per 1.5 row-meters. Final tonnage
of plant-cane stalks per ha was reduced by 5.9 metric tons per wireworm per 1.5 row-m
(r = .92, F = 81.8, P > F = .0001), or by 3.8% per wireworm per 1.5 row-m (r = .92,
F = 72.69, P > F = .0001). No reductions were observed in sucrose levels in juice
from plots infested by wireworms. During the ratoon crop, plots that had been infested
by 8 or 12 wireworms per 1.5 row-meters during the plant cane crop had significantly
fewer shoots. General observations indicated that these stand reductions were strictly
a result of wireworm damage that occurred during the plant-cane crop. Final tonnage


Plant-cane crop
x No. First-ratoon crop
t No. stalks per ha
wireworms after planting i Metric R % i No. x Metric
per 1.5 tons of cane yield ratoon stalks tons of cane
row-meters 8 Wks 12 Wks per ha 96 sugar per ha2 per ha

0 22,656a 34,297a 153.0a 10.4a 94,956a 89.8a
4 15,709b 22,781b 128.2a 10.5a 89,573ab 76.6ab
8 8,887c 12,830c 101.6b 10.6a 64,839bc 72.4b
12 6,008c 7,072c 83.2b 10.7a 51,771c 53.1c

'Means in the same column followed by the same letter are not significantly different using the Ryan-Einot-Gab-
riel-Welsch multiple range test. F-values (P > F) for the six ANOVA's (from left to right) were 36.7 (.0001), 59.7
(.0001), 21.1 (.0002), 0.7 (.5964), 14.6 (.0003), and 12.2 (.0016), with 9 error d.f. for each.
'Ratoon stand determined August 17, 1989, 23 weeks after the plant cane was harvested.

Florida Entomologist 73(2)

June, 1990

of the first-ratoon cane was significantly reduced in plots that had been infested during
plant cane at 8 and at 12 wireworms per 1.5 row-m. Tonnage of the ratoon cane was
reduced by 2.9 metric tons per ha (about 3.2%) for each wireworm present per 1.5
row-m during early plant cane growth (r = .86, F = 38.3, P > F = .0001). The overall
drop in tonnage between the plant and ratoon crops was larger than normal for CL
61-620 due to freezes that occurred near the end of the ratoon crop.
The data from these studies along with those from an earlier study (Hall 1985)
indicated that significant stand reductions during early plant-cane growth may occur at
infestation levels of fewer than four M. communis per 1.5 row-meters. Wireworms may
cause larger stand reductions in varieties that emerge slowly and/or that produce fewer
stalks. Stand reductions per wireworm per 1.5 row-m may range from around 6 to 8%.
Tonnage reductions caused by M. communis may range from around 3 to 4% per
wireworm per 1.5 row-m; in the 1988 large plot study, yield reductions were attributed
to early stand losses caused by wireworms because tonnage produced per hectare was
highly correlated with the stand of shoots at 12 weeks after planting (r = .96, F =
145.4, P > F = .0001). Although wireworm infestations did not reduce sucrose levels
in juice, the amount of sugar produced per hectare was reduced due to the tonnage
reductions. For example, a 3.5% tonnage reduction would represent a 3.5% reduction
in sugar production. The results of this research showed that stand reductions caused
by wireworms during plant cane will reduce the profitability of ratooning.


Ron P. DeStefano, chief chemist at U. S. Sugar Corp., conducted sucrose analyses.


BOURNE, B. A. 1968. Important key which aided greatly the sugar cane research work
in Florida. Sugar Jour. 30(8): 11-13 and 30(9): 29.
CHERRY, R. H., AND D. G. HALL. 1986. Flight activity of Melanotus communis
(Coleoptera: Elateridae) in Florida sugarcane fields. J. Econ. Entomol. 79(3):
GIFFORD, J. R. 1964. A brief review of sugarcane insect research in Florida. 1960-
1964. Proc. Soil and Crop. Sci. Soc. Florida. 24: 449-453.
HALL, D. G. 1985. Damage by the corn wireworm, Melanotus communis (Gyll.), to
plant cane during germination and early growth. J. Amer. Soc. Sugar Cane
Techn. 4: 13-17.
- 1988. Insects and mites associated with sugarcane in Florida. Florida Entomol.
71(2): 138-150.
INGRAM, J. W., H. A. JAYNES, AND R. N. LOBDELL. 1938. Sugarcane pests in
Florida. Proc. Internat'l. Soc. Sugar Cane Techn. 6: 89-98.
TIER. 1951. Pests of sugarcane and their control. U.S.D.A. Cir. 878. 38 pp.
SAS INSTITUTE. 1985. SAS user's guide: statistics, version 5 ed. SAS Institute,
Cary, N.C.
WARD, R. H., AND A. J. KEASTER. 1977. Wireworm baiting: use of solar energy to
enhance early detection of Melanotus depressus, M. verberans, and Aeolus mel-
lillus in midwest cornfields. J. Econ. Entomol. 70(4): 403-406.


Atkinson et al.: Annotated Checklist of Florida Cockroaches 303


Department of Entomology and Nematology,
University of Florida,
Gainesville, Florida 32611-0540

Insects Affecting Man and Animals Research Laboratory, USDA-ARS,
1600 SW 23rd Dr., Gainesville, Florida 32608


Synonymy, distribution, and ecological data are summarized for 38 species of cock-
roaches in 24 genera in 4 families which occur in Florida. We include information on
other species known from nearby areas which may also be collected in the state. Thir-
teen of these species, including most of the important pests, have been introduced from
Africa (7), the Neotropics (3), and Asia (3). Most exotic species appear to be dependent
on human disturbance and only two, Pycnoscelis surinamensis (L.) and Periplaneta
australasiae (F.), are commonly found in natural communities away from human distur-
bance. Blattella germanica (L.) and Supella longipalpa (F.), both introduced, are
strictly domiciliary. Most of the native species (15) have neotropical distributions and
are not found north of Florida. Four species, Chorisoneura texensis Saussure &
Zehntner, Euthlastoblatta gemma (Hebard), Eurycotis floridana (Walker), and Is-
chnoptera deropeltiformis (Brunner), are restricted to the southeastern U.S. and are
the northernmost representatives of neotropical genera. Five species of Parcoblatta,
distributed widely in the Southeast, reach their southern limits in Florida. There is one
endemic species, Arenivaga floridensis Caudell, restricted to sandy areas of central


Se resume sinonimia, distribuci6n, y datos ecol6gicos para 38 species de cucarachas
en 24 generos en 4 families que ocurren en la Florida. Incluimos informaci6n sobre otras
species conocidas de areas cercanas las cuales pueden colectarse en el estado. Trece
de 6stas, incluyendo la mayor parte de las plagas importantes, se han introducido de
Africa (7), del Neotr6pico (3), y de Asia (3). La mayoria de las species ex6ticas parecen
defender del disturbio human y solo dos, Pycnoscelis surnamensis (L.) y Periplaneta
australasiae (F.), se encuentran cominmente en comunidades naturales lejos de distur-
bio human. Blattella germanica (L.) y Supella longipalpa (F.), ambas introducidas,
son estrictamente domiciliarias. La mayoria de las species nativas (15) tienen dis-
tribuciones neotropicales y no se encuentran al norte de Florida. Cuatro species,
Chorisoneura texensis Saussure & Zehntner, Euthlastoblatta gemma (Hebard),
Eurycotis floridana (Walker), y Ischnoptera deropeltiformis (Brunner), se limitan al
sureste de los EE. UU. y son las representantes mas septentrionales de g6neros neot-
ropicales. Cinco species de Parcoblatta, distribuidas ampliamente en el sureste, alcan-
zan sus limits australes en Florida. Hay una especie endemica, Arenivagafloridensis
Caudell, restringida a areas arenosas de la Florida central.

The Asian cockroach, Blattella asahinai Mizukubo (Roth 1986), is probably the most
recent addition to the rich cockroach fauna of Florida. Unlike its close relative, the

304 Florida Entomologist 73(2) June, 1990

German cockroach, B. germanica (L.), the Asian cockroach is essentially an outdoor
species, only entering homes occasionally (Brenner et al. 1988). We began reviewing
the literature on species already present in the state, both native and introduced, for
purposes of identification of this new exotic and similar species and as preparation for
studies on its potential interactions with other outdoor species. We were struck by the
general paucity of up-to-date information on the cockroach fauna, exclusive of the well-
documented domestic and peridomestic pest species.
The most recent comprehensive taxonomic treatment of the cockroaches of the con-
tinental United States was that of Hebard (1917) and included 46 species from the U.S.
More recent general works include keys by Blatchley (1920) and Heifer (1963) and a list
of names by Pratt (1988) which added virtually no new biological or distributional infor-
mation. Pratt (1988) included 66 species for the U.S. Some of this increase was due to
introduction and establishment of exotic species. More thorough collection in southern
Florida and the Southwest also resulted in descriptions of new species and detection of
other species which were previously known from neighboring areas and probably do not
represent introductions.
This checklist was prepared to summarize synonymy, distributions, and ecology of
all species of cockroaches known from Florida and adjacent areas. We were particularly
interested in analyzing and comparing distribution and ecology of native and introduced
species, with emphasis on their pest status.


The majority of the information presented here is summarized from an exhaustive
literature review. Specimens deposited in the Florida State Collection of Arthropods
were studied by the senior author for additional distributional information. We have
also included information from an unpublished inventory of the entomofauna of the
Archbold Biological Station, Lake Placid, Highlands Co. (M. A. Deyrup, personal com-
munication). The bulk of the available information on distributions in Florida is based
on collections made prior to 1920 by Hebard, Rehn, Davis, and Blatchley (Blatchley
1920, Davis 1914, 1915, Hebard 1916, Rehn & Hebard 1904, 1905, 1907, 1910, 1914a,b).
Subsequent information has been scattered among ecological studies (e.g. Friauf 1953,
Peck & Beninger 1989) or miscellaneous taxonomic notes.
Hebard's (1917) monograph was taken as our starting point and no attempt was
made to review prior literature. Previous publications, including those of Hebard him-
self, contained many misidentifications and other incorrect usage of names. Synonymies
and overall distributions were summarized from Princis' treatment of the Blattaria in
the Orthopterorum Catalogus (1962, 1963, 1964, 1965, 1966, 1967, 1969, 1971). These
were modified by more recent treatments when applicable.
The order of superfamilies and families follows McKittrick (1964), and differs consid-
erably from that used by Princis. Species are listed alphabetically within genera, and
genera alphabetically within families.
We include complete synonymy for each species. Genera in which a given species
name has been published are included in parentheses if these combinations have not
been used in the North American literature. Novel combinations or important misiden-
tifications in North American usage are indicated. The nomenclatorial usage of impor-
tant taxonomic and general references is indicated but our listings are not exhaustive
in this regard. We have listed in the synonymy all references in the North American
literature since 1917 that deal with taxonomy, identification and description of life
stages, and ecology, particularly with reference to outdoor habitats and distribution
with respect to structures. Not all of these are cited in the text of the article, but the
full bibliographic citations are included among the references cited as an aid to readers.

Atkinson et al.: Annotated Checklist of Florida Cockroaches 305

All common names which have been used in the literature are listed also. Most were
invented by Blatchley (1920), Helfer (1963), or Pratt (1988) and do not reflect common
use. Common names that are on the approved list of the Entomological Society of
America (Werner 1982) are indicated with an asterisk. The name "palmettobug" is
indiscriminately used for any large cockroach, principally species of Periplaneta and
Eurycotis (Gurney & Walker 1976).
Distributional information includes a summary of worldwide distribution, list of
states and provinces in the U.S. and Canada, and counties in Florida. We have used
numbered superscripts in the distribution lists to indicate literature citations to save
space and to reduce distraction. An asterisk indicates a previously unreported locality.
Biological and ecological information is summarized where available with emphasis on
relative abundance in and about structures, disturbed areas, and natural habitats.
Biological information on the important domestic and peridomestic pest species is co-
vered more adequately in references such as Cornwell (1968) and is not repeated.


Four additional species have been reported from Florida that were not on Pratt's
list (1988): Blaberus discoidalis Serville (Roth 1969), Myrmecoblatta wheeler Hebard
(Deyrup & Fisk 1984), Neoblattella detersa (Walker) (Peck & Beninger 1989), and
Symploce morse (Peck & Beninger 1989). Chorisoneura parishi Rehn, apparently es-
tablished in Miami, is reported from Florida and the U.S. for the first time here. Pratt
(1988) and Princis (1969) listed Euthlastoblatta diaphana (F.) for the southeastern
U.S., but it does not occur in the U.S. They apparently overlooked Hebard's comments
(1917) on misidentifications of E. gemma Hebard under this name. Taking these addi-
tions and deletions into account, there are at least 69 species of cockroaches in 31 genera
in 5 families known from the continental U.S.
Cockroaches generally are associated with litter and decaying wood in forest ecosys-
tems. They reach their greatest taxonomic diversity in the humid tropics and subtropics
(Schal et al. 1984). Not surprisingly, Florida, with its mild humid climate and variety
of subtropical and temperate forest ecosystems has the richest cockroach fauna of any
part of the U.S. Currently 38 of the 69 species known from the U.S. are found in
Florida. Thirteen of these have been introduced. More native and introduced species
likely would be detected with more field work by knowledgeable collectors.
Distributions of the 25 native species known from Florida are summarized in Table
1. A very strong tropical bias is immediately evident. Most of the genera (15 of 7) are
neotropical and 15 of the 25 species are found in the Caribbean and/or Mesoamerica,
reaching their northern limits in Florida. Several of these neotropical species are known
only from the Keys and have been considered by previous authors to be introduced
(e.g., Hebard 1917:260 re Blaberus craniifer Burmeister, Phoetalia pallida (Brunner),
Holocompsa nitidula (F.)). In the absence of any convincing evidence to the contrary
(e.g., Panchlora nivea (L.)), we take the conservative approach of treating most of
these as natives (i.e., not introduced by man).
Chorisoneura texensis Saussure & Zehntner, Euthlastoblatta gemma (Hebard),
Eurycotis floridana (Walker), and Ischnoptera deropeltiformis (Brunner), are re-
stricted to the southeastern U.S., but are the northernmost representatives of neotrop-
ical genera.
Five species of Parcoblatta, a nearctic genus, reach their southernmost limits in
Florida, only 1 of which is found into southern Florida. Two other species of Parcoblatta,
P. bolliana (Saussure & Zehntner) and P. pensylvanica (DeGeer), are known from
adjoining counties in southern Georgia and probably occur in northern Florida.

Florida Entomologist 73(2)


Number of Species

Family Genus Genus dist. FL SFL CFL NFL NTRP GC EUS

Eurycotis Neotropical 2 2 1 1 1 1 0
Arenivaga SW Nearctic 1 1 1 -
Compsodes Neotropical 2 2 1 -
Holocompsa Neotropical 1 1 1 -
Myrmecoblatta Neotropical 1 ? 1 ? 1 ? -
Cariblatta Neotropical 2 2 2 1 2 1 1
Chorisoneura Neotropical 1 1 1 1 1 1
Euthlastoblatta Neotropical 1 1 1 1 1 -
Ischnoptera Neotropical 1 1 1 1 1 1
Latiblattella Neotropical 1 1 1 1 -
Neoblattella Neotropical 1 1 1 -
Parcoblatta Nearctic 5 1 1 5 5 5
Plectoptera Neotropical 1 1 1 -
Symploce Neotrop/Afr. 1 1 1 -
Blaberus Neotropical 2 2 1 -
Hemiblaberus Neotropical 1 1 1 -
Phoetalia Neotropical 1 1 1 -

Totals 25 19 10 11 15 10 8

The only endemic cockroach species known from Florida, Arenivaga floridensis
Caudell, is restricted to sandy areas of central Florida and is the only eastern represen-
tative of the genus. The remaining species of Arenivaga are known from temperate
desert and semiarid areas of southwestern North America (Hebard 1920).
The only native species which might be considered a pest is the Florida woods roach
or palmetto bug, Eurycotis floridana, which breeds around houses in shaded, moist
environments, and occasionally enters buildings. All our major pest species are exotics.
Information on the 13 introduced species is summarized in Table 2. Most species
originated in Africa (7), followed by Asia (3) and the Neotropics (3). Infestations of
Nauphotea cinerea Olivier) were reported from feed mills in the Tampa area in the
early 1950's (Gresham 1952). Since that time there have been no further notices of its
activities or indication that it has spread, suggesting that the infestation did not persist.
The only species which have successfully invaded naturally occurring communities in
the state are the Australian cockroach, Periplaneta australasiae, and the Surinam
cockroach, Pycnoscelus surinamensis (L.). The Cuban cockroach, Panchlora nivea
(L.), is common in wooded lots within the urbanized area of Gainesville, FL, and may
also be established in similar forested areas further removed from human activity. Most
of the other species appear to depend on human disturbance to persist, with the possible
exception of some of other species of Periplaneta (e.g., fuliginosa). Blattella germanica
(L.) and Supella longipalpa (F.) represent the extreme of dependency and could be
considered obligate domiciliary species in Florida. While individuals of these species
may be found out of doors, breeding populations either do not occur or do not persist.

June, 1990

Atkinson et al.: Annotated Checklist of Florida Cockroaches 307


Family/Species D P N Origin Distribution in U.S.

Periplaneta + + Africa throughout U.S. as urban-
americana domiciliary, outdoors in Southeast
P. australasiae + + + Africa outdoors in central & southern
Florida, further northwards as
P. brunnea + Africa southeastern U.S. to southern Fla.
P. fuliginosa + ? Africa southeastern U.S., not in
peninsular Fla.
Blattella asahinai ? + ? Asia peninsular Fla.
B. germanica + Asia entire U.S., obligate domiciliary
Chorisoneura + Caribbean southern Fla.
Supella longipalpa + Africa entire U.S., obligate domiciliary
Symploce pallens + Africa Key West
Epilampra maya ? ? Mesoamerica Arcadia
Nauphoeta cinerea + ? Africa Tampa
Panchlora nivea + ? Neotropics peninsular Fla., Gulf Coast
Pycnoscelis + + trop. Asia outdoors in Fla., Gulf Coast,
surinamensis further north in greenhouses, etc.



1. Blatta orientalis L.
Oriental cockroach*

Blatta orientalis Linnaeus 1758:424 Syst. Nat., X ed.; Hebard 1917:173 (tax, fig);
Blatchley 1920:94 (tax, fig); Hebard 1943:271 (tax); Rehn 1945:266 (dispersal); Froes-
chner 1954:180 (tax); Helfer 1963:50 (key, fig); Princis 1965:475 (tax); Cornwell 1968
(biol, ecol, econ status); Dakin & Hays 1970:12 (tax); Pratt 1988:883 (tax). (Kakerlac,
Periplaneta, Stylopyga).
Blatta secunda Schaeffer 1769:155 Icones Ins. circa Ratisbonam indigen.
Blatta tertia Schaeffer 1769:155 Icones Ins. circa Ratisbonam indigen.
Blatta culinaris DeGeer 1773:530 Mem. Hist. d. Ins.
Blattaferruginea Thunberg 1810:187 Vetenskapsakad. nya Handl.
Blatta europaea 1846:30 Saros magye helyirata.
Kakerlac castanea Blanchard 1851:18 Hist. Fis. Polit. Chile.
Pulex imperator Westwood 1858:70 Trans. Entomol. Soc. Lond.
Blatta badia Saussure 1863:150 Mem. Soc. Geneve.
Kakerlac pallipes Philippi 1863:222 Z. Ges. Naturwiss. (Periplaneta, Stylopyga).
Kakerlac platystetho Philippi 1863:222 Z. ges. Naturwiss. (Periplaneta).
Blatta germanica (not germanica L.) Burr 1904:120 Entomol. Rec.
Stylopyga orientalis var. gracilis Adelung 1910:337 Soc. Entomol. Ross.
DISTRIBUTION: Cosmopolitan in temperate areas. AL5,12 AZ'2, CA", CO12, CT12, IA8.1
IL15, IN', KS14, MIP, MN2, MO12, NC12, NE12, NJ12, NM12, PA'2, TN12, TX126, WI12.

308 Florida Entomologist 73(2) June, 1990

Blatchley (1920) reported this species from Florida (Miami and West Palm Beach),
however these records are undoubtedly based on adventive collections because there is
no evidence that breeding populations B. orientalis occur anywhere in Florida or on the
Southeastern Coastal Plain. We are aware of unconfirmed anecdotal reports of this
species in the Jacksonville area, but have seen no specimens. Given its abundance and
pest status in cooler parts of the U.S., it is likely to be introduced periodically and
specimens may be found occasionally.
ECOLOGY: This species occurs commonly in and about houses and other structures over
most of the temperate U.S. It apparently does not occur in natural communities away
from human disturbance in the parts of the country where it is found.

2. Eurycotis floridana (Walker)
Florida cockroach, Florida woods roach, palmettobug

Perplaneta floridana Walker 1868:135 Cat. Blatt. British Mus. (Pelmatosilpha).
Eurycotis floridana: Hebard 1917:166 (tax, fig); Blatchley 1920:97 (tax, fig); Friauf
1953:122 (ecol); Helfer 1963:49 (fig male, female); Princis 1966:553 (tax); Dakin &
Hays 1970:13 (tax); Gurney & Walker 1976:824 (biol., ecol.); Hagenbuch et al.
1988:378 (ecol); Brenner 1988:583 (ecol); Pratt 1988:883; Patterson & Koehler 1989:39
(ecol, cont); Peck & Beninger 1989:614.
Periplaneta semipicta Walker 1868:141 Cat. Blatt. British Mus.
Platyzosteria ingens Scudder 1877:92 Proc. Boston Soc. Nat. Hist.
Platyzosteria sabalianus Scudder 1877:93 Proc. Boston Soc. Nat. Hist.
DISTRIBUTION: Peninsular Florida, lower Gulf and Atlantic coasts: AL5, FL'2," GA12,
MS12. In Florida: Alachua", Clay12, Dade'2, Duval12, Escambia12, Levy12, Highlands
(M.A. Deyrup pers. comm. 1989), Hillsborough12, Polk12, Putnam8, Charlotte'", Citrus12,
Collier'2, Broward'1, Dade12, Monroe2,20.
ECOLOGY: Common in native vegetation as well as near human habitation, this species
will enter houses occasionally, but does not commonly breed indoors. This species re-
leases an oily, irritating liquid with a strong odor when disturbed and is sometimes
referred to as the "stinking cockroach" or "Florida stinkbug" (Blatchley 1920, Gurney
& Walker 1976). Breeding has been detected in attics (R. J. Brenner, pers. comm.). It
is found in many native communities throughout Florida (Peck & Beninger 1989, Friauf

3. Eurycotis lixa Rehn
Hustler cockroach

Eurycotis lixa Rehn 1930:45 Trans. Amer. Entomol. Soc.; Gurney 1959:75 (occurrence
in Florida, tax, fig female); Helfer 1963:49 (key); Princis 1965:550 (tax); Pratt
1988:883 (checklist).
DISTRIBUTION: Jamaica"'. Florida: Monroe (Key West)10. Gurney (1959) considered this
to be an introduced species, but it is also present in the West Indies and its "immigrant"
status is questioned.

4. Periplaneta americana (L.)
American cockroach*

Blatta americana L. 1758:424 Syst. Nat., 10th ed. (Kakerlac)
Periplaneta americana: Hebard 1917:176 (tax, fig), Blatchley 1920:99 (tax, fig); Hebard
1943:269 (tax); Rehn 1945:269 (dispersal); Friauf 1953:122 (ecol); Froeschner 1954:181
(tax); Helfer 1963:51 (key, fig); Princis 1965:405 (tax); Cornwell 1968 (gen info, biol,
econ status); Dakin & Hays 1970:12 (tax); Powell & Robinson 1980:216 (1st instar

Atkinson et al.: Annotated Checklist of Florida Cockroaches 309

nymph); Hagenbuch et al. 1988:378 (ecol); Brenner 1988:583 (ecol); Pratt 1988:883
Blattaferrugineo-fusca Gronovius 1764:174 Zoophylac. Gronov. 2.
Blatta kakkerlac DeGeer 1773:535 Mem. l'Hist. Ins.
Blatta aurelianensis Fourcroy 1785:177 Entomol. Parisiensis 1.
Blatta siccifolia Stoll 1813:5 Repres. exact, coloree nature d. Spectres ..
Blatta heros Eschscholtz 1822:83 Entomographica Erste Lief.
Periplaneta stolida Walker 1868:129 Cat. Blatt. Brit. Mus.
Perilpaneta americana colorata Rehn 1901:220 Trans. Amer. Entomol. Soc.
DISTRIBUTION: Cosmopolitan, northwards to New York City in eastern U.S.: AL12,5,
AZ12, CA12, FL121" GA12, IA', IL16, LA12, MI3,MS12, NC12, NY12, PA12, TX12. In
Florida: Alachua", Highlands (M.A. Deyrup pers. comm. 1989); Polk'1, Monroe (incl.
Keys)"1, Putnam". This species is probably found in all Florida counties and all of the
U.S. states. It is generally considered to be widely distributed and we have no reason
to doubt this. Nonetheless it is curious that there is relatively little documentation of
its current distribution in the U.S. In the southern part of its range this species may
breed outdoors.
ECOLOGY: In an ecological study in northeastern Florida this species was found only
around buildings, not in native habitats (Friauf 1953). A similar situation occurs at the
Archbold Biological Station in sandhill communities in central Florida (M.A. Deyrup
pers. comm. 1989). It is a common peridomestic species in northern Florida (Hagenbuch
et al. 1988).

5. Periplaneta australasiae (F.)
Australian cockroach*

Blatta australasiae Fabricius 1775:217 Syst. Entomol.
Periplaneta australasiae: Hebard 1917:185 (tax, fig); Blatchley 1920:101 (tax, fig); Rehn
1945:269 (dispersal); Friauf 1953:122 (ecol); Froeschner 1954:182 (tax); Helfer 1963:51
(key, fig); Princis 1965:447 (tax); Lawson 1967:269 (ecol, dist); Cornwell 1968 (gen
info, biol, econ status); Powell & Robinson 1980:222 (1st instar nymph); Hagenbuch
et al. 1988:378 (ecol); Pratt 1988:883 (checklist); Patterson & Koehler 1989:39 (ecol,
Blatta domingensis Palisot de Beauvois 1805:182 Ins. Rec. Afr. et Amer.
Blatta aurantiaca Stoll 1813:5 Represent. exact. coloree d'apres des Spectres ...
Periplaneta zonata Haan 1842:49 Verhand. natural. Gescheid. Nederl. over. Bezitt.
Periplaneta repanda Walker 1868:125 Cat. Blatt. Brit. Mus.
Periplaneta subcincta Walker 1868:126 Cat. Blatt. Brit. Mus.
Periplaneta inclusa Walker 1868:126 Cat. Blatt. Brit. Mus.
Polyzosteria subornata Walker 1871:35 Cat. Derm. Salt. Brit. Mus., Suppl. Blatt.
Periplaneta emittens Walker 1871:37 Cat. Derm. Salt. Brit. Mus., Suppl. Blatt.
DISTRIBUTION: Florida: Alachua", Charlotte12, Clay', Collier12, Dade2, Highlands
(M.A. Deyrup pers. comm. 1989), Hillsborough',18, Levy12, Monroe (incl. Keys)'2,
Orange"2, Pinellas', Seminole'. This species is occasionally found indoors in many parts
of the U.S. but apparently breeds outdoors only in central and southern Florida.
ECOLOGY: In an ecological study in northeastern Florida this species was found only
around buildings, not in native habitats (Friauf 1953). At the Archbold Biological Station
in sandhill communities in central Florida this species is found in and near buildings and
is also common in native vegetation (M. A. Deyrup pers. comm. 1989). It is a common
peridomestic species in north-central Florida (Hagenbuch et al. 1988) and occurs in
treeholes, palm trees, and voids in block walls used in landscaping (R. J. Brenner, pers.

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