Title: Florida Entomologist
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098813/00072
 Material Information
Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1989
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
 Record Information
Bibliographic ID: UF00098813
Volume ID: VID00072
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 72, No. 3 September, 1989


Preface .................................................................................................. 391
KOEHLER, C. S.-Prospects ofImplementation ofIPM Programs for Ornamental
P lants .......................................................................................... 391
in Sampling in Ornamentals ............................................. ....... 394
REICHELDERFER-Economic and Aesthetic Injury Levels and Thresholds
for Insect Pests of Ornamental Plants .............................................. 403
OSBORNE, L. S., AND R. D. OETTING-Biological Control of Pests Attacking
Greenhouse Grown Ornamentals ...................................................... 408
NIELSEN, D. G.-Exploiting Natural Resistance as a Management Tactic for
Landscape Plants ........................................................................... 413

COOK, C.-Philogenia redunca, A New Damselfly From Ecuador (Odonata:
M egapodagrionidae) ........................................................................ 419
DONNELLY, T. W.-A New Species of Philogenia From Honduras (Odonata:
M egapodagrionidae) ........................................................................ 425
DONNELLY, T. W.-Three New Species of Epigomphus From Belize and Mexico
(Odonata: Gomphidae) ................................................................. 428
DONNELLY, T. W.-Protoneura sulfurata, A New Species of Damselfly From
Costa Rica, With Notes on the Circum Caribbean Species of the Genus
(Odonata: Protoneuridae) ................................................................ 436
SPENCER, K. A., AND D. HAVRANEK-A New Species of Agromyzidae (Diptera)
From Venezuela ............................................................................. 441
PENA, J. E., R. J. GAGNE, AND R. DUNCAN-Biology and Characterization of
Prodiplosis longifilia (Diptera: Cecidomyiidae) on Lime in Florida ....... 444
STANGE, L. A.-Review of the New World Dimarini, With the Description of a
New Genus From Peru (Neuroptera: Myrmeleontidae) ........................ 450
O'BRIEN, C. W., AND H. R. PAJNI-Two Indian Bagous Weevils (Coleoptera:
Curculionidae), Tuber Feeders of Hydrilla verticillata (Hydrocharitaceae),
One a Potential Biocontrol Agent in Florida ...................................... 462
HOWARD, F. W., AND M. A. SOLIS-Distribution, Life History, and Host Plant
Relationships of Mahogany Webworm, Macalla thyrsisalis (Lepidoptera:
Pyralidae) ..................................................................................... 469
GIBLIN-DAVIS, R. M., K. GERBER, AND R. GRIFFITH-Laboratory Rearing of
Rhyncophorus cruentatus and R. palmarum (Coleoptera: Curculionidae)
..... ......................... .......................... ........................ ......... ...... 480
PROKOPY, R. J., AND D. R. PAPAJ-Can Ovipositing Rhagoletis pomonella
Females (Diptera: Tephritidae) Learn to Discriminate Among Different
Ripeness Stages of the Same Host Biotype? ....................................... 489
HILBURN, D. J.-A Non-Migratory, Non-Diapausing Population of the Monarch
Butterfly, Danaus plexippus (Lepidoptera: Danaidae), in Bermuda ........ 494

Continued on Back Cover

Published by The Florida Entomological Society


President ........................................ ... ................. J. E Eger
President-Elect ........................ .................................. 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
C. O. Calkins
F. Bennett
J. E. Pefia
N. Hinkle
M. F. Antolin
J. R. McLaughlin


Editor ........................................ .. ........... ................... J. R. McLaughlin

Associate Editors

Arshad Ali
John B. Heppner
John Sivinski
Willis W. Wirth

Carl S. Barfield
Michael D. Hubbard
Omelio Sosa, Jr.

Ronald H. Cherry
Lance S. Osborne
Howard V. Weems, Jr.

Business 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.
Membership in the Florida Entomological Society, including subscription to Florida
Entomologist, is $25 per year for regular membership and $10 per year for students.
Inquires regarding membership, subscriptions, and page charges should be addres-
sed to the Business 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 Winter Haven, FL.
Manuscripts from all areas of the discipline of entomology are accepted for consider-
ation. At least one author must be a member of the Florida Entomological Society.
Please consult "Instructions to Authors" on the inside back cover. Submit the original
manuscript, original figures and tables, and 3 copies of the entire paper. Include an
abstract in Spanish, if possible. Upon receipt, a manuscript is acknowledged by the
Editor and assigned to an Associate Editor who sends it out for review by at least 3
knowledgeable peers. Reviewers are sought with regard only for their expertise; Soci-
ety membership plays no role in their selection. Page charges are assessed for printed
Manuscripts and other editorial matter should be sent to the Editor, JOHN R.
MCLAUGHLIN, 4628 NW 40th Street, Gainesville, FL 32606.

This issue mailed September 15, 1989




During the last 30 years there has been a phenomenal population shift in the United
States. Today, some 80% of people live, work, and recreate in urban/suburban settings.
A recent Gallup poll ranked gardening and landscaping activities first nationwide among
outdoor pursuits undertaken for recreation, health, and economy. The popularity of this
national pastime has created a growing demand for pest free ornamental plants for
landscape and interior use which now supports a thriving production and retail industry.
In the last 10 years, newly adopted pesticide use laws, and fear of litigation from
pesticide misuse in urban settings, have become an increasing concern. In reaction to
this situation, commercial and private pesticide applicators in the green industries have
begun adopting the IPM approach for ornamental plant pest management. In turn, this
has stimulated a budding, but still underfunded, core of dedicated university research
and extension personnel. Their goal is to provide technical support to a growing group
of IPM practitioners.
The papers which follow were presented in the first formal conference on ornamental
plant IPM presented at a meeting of the Entomological Society of America. They reflect
the current status of ornamental plant IPM theory and practice. They clearly show that
much work remains to be done.

Department of Entomology
University of Maryland
College Park, MD 20742


Cooperative Extension, University of California
Berkeley, California 94720


Public concern over perceived hazards of pesticides, increasing costs of conventional
pest control, and legislation such as the Endangered Species Act all serve in their own
way to advance the prospects for implementation of IPM programs for ornamentals.
Constraints on pesticide use enacted by legislatures, however, seldom bring forth new
funding for IPM research. Beyond funding shortfalls for such research, the prospects
for IPM advancement are further diminished by inadequate numbers of ornamentals
research and practitioner personnel. The costs for providing IPM services to consumers
are no less than those required to provide conventional spraying services.


Inquietud pfiblica sobre el peligro de pesticides, el aumento del costo del control
conventional de plagas, y legislaci6n tal como el Acta de Especies en Peligro, todas

392 Florida Entomologist 72(3) September, 1989

sirven, en su propia manera, para avanzar el prospect de la implementaci6n de prog-
ramas de control integrado de plagas en plants ornamentales. Restricciones en el uso
de pesticides han sido legisladas, sin embargo, pocas veces han producido un aumento
de fondos econ6micos para investigaciones en programs de control integrado. Aparte
de la escazes de fondos para tales investigaciones, los prospects de advance en los
programs de control integrado se disminuyen tambi6n por el nfimero inadecuado de
personas envueltas en investigaciones de plants ornamentales. El costo de dar servicio
al consumidor en programs de control integrado no son menos que auellos necesarios
para proveer servicios convencionales de rocios.

The objectives of this report are to look at the forces that drive IPM forward, and
also at those factors that serve to impede it. Reference will be made both to production
and landscape ornamentals.
Nearly everyone will agree that the public's concern about the perceived human
health risks of pesticides, and their degradation of the environment, is a strong force
driving IPM forward. The media's willingness to report and interpret the significance
of every pesticide incident is creating a public which is becoming increasingly receptive
to IPM and its often accompanying reduction in pesticide usage. Public opinion often
becomes translated into legislation which will augment IPM acceptance.
There are many reports of impressive reductions in pesticide use through IPM in
the landscape. These vary between 50 and 95 percent, yet I remain skeptical about a
reported dramatic reduction in pesticide usage unless it is accompanied by an evaluation
of the appearance of the target vegetation before and after the IPM program.
On the subject of legislation, all signs suggest that we are going to see fewer of the
moderately to highly toxic insecticides being used in the landscape, at least as sprays.
The posting of treated properties, and the requirement to notify neighbors before the
application, keep surfacing in legislative chambers. Of newest interest is the exact way
in which the EPA, and state regulatory agencies, will enforce the Endangered Species
Act, as it relates to pesticides. One California official noted recently that it is no longer
a question of whether the Act will be enforced, only when and how. According to a
report by the California Nature Conservancy, about 10 percent of California's native
mammals, 17 percent of native reptiles and amphibians, and 27 percent of freshwater
fish are listed by the state or federal governments as threatened or endangered species
(California Nature Conservancy 1987). If species waiting to be listed were added, fully
a third of California's mammals, a quarter of the birds, a third of the reptiles and
amphibians and 40 percent of the freshwater fish would be classified as imperiled. The
geographic ranges of these species occupy huge acreages in California. Agriculture is
most concerned about the Endangered Species Act just now, but whatever applies to
agriculture will also apply to the landscape not planted to agricultural crops. Rigid
enforcement of the Endangered Species Act can only serve to drive landscape IPM
IPM in some cases is driven by economics. This is far more true for production
ornamentals than for those in the landscape. Parrella & Jones (1987) note the
documented or suspected resistance in the greenhouse of the leafminer Liriomyza
trifolii (Burgess), beet armyworm, greenhouse whitefly, twospotted spider mite, west-
ern flower thrips, and green peach aphid. When growers are confronted with severe
resistance problems, and, when faced with the non-availability of new and effective
insecticides, they become extremely receptive to alternative methods of pest control.
In a few cases IPM has been demonstrated to offer such an alternative.
Competition from foreign-produced flowers is another economic force which may
tend to drive greenhouse crop growers toward IPM. This is particularly true if domestic

_ __ __~

Koehler: Symposiumr-IPM for Ornamentals 393

costs of producing flowers continue to escalate because of the increasing costs of pes-
ticides or other traditional forms of pest control.
Economics as a driving force for IPM is not so apparent in landscape ornamentals.
Here, we have relatively little pesticide resistance that has been documented or sus-
pected, and for most of the important, and even minor, pests, effective pesticides are
available. Resistance costs, therefore, are not an issue.
Recent reports by Smith & Raupp (1985) and Smith & Gill (1987) compared the
operating costs of a traditional cover spray vs. an IPM program in a planned community
complex in Maryland. Scouts were hired, then trained by Cooperative Extension per-
sonnel to monitor, at 2 to 3 week intervals, the landscape vegetation on nearly 500
acres. In the first year of the program, plants identified as needing spraying were
actually treated by a commercial arborist. In the second and final year, the spot spraying
of low vegetation was done directly by the monitoring personnel, but an outside
applicator was called in to treat the taller plants on an as-needed basis.
While the average saving, through IPM, was reported to be 22 percent-with no
sacrifice in the appearance of plants in the IPM program-I question whether all the
appropriate costs for IPM and the conventional program really were calculated. For
example, the training of IPM scouts was done by Cooperative Extension, a subsidized
cost. Certain plant problem diagnoses were referred to University of Maryland
Specialists, another subsidized cost. Almost certainly the liability insurance, and a cer-
tain profit, were costs in the traditional cover spray program, but not in the IPM pilot
program. In my opinion we'll never sell IPM in landscape ornamentals on the basis of
cost saving. Most of the private sector firms I am acquainted with, who offer an IPM
alternative to their landscape customers, charge the same for either the conventional
or the IPM service.
The impediments to implementation of ornamentals IPM are varied. Funding levels
for IPM research in neither production nor landscape ornamentals are sufficient for the
magnitude of the job to be done. Until or unless this situation changes for the better,
progress in implementing IPM will be seriously impeded.
Much of the funding shortfall, especially in landscape ornamentals, is related to the
fact that the principal benefactor of IPM has no heartbeat and therefore no pulse.
Unlike the cotton, corn, or wine interests, homeowners or many others who gain from
IPM are not organized to beat on the legislative doors at Albany, Annapolis, or Sac-
ramento for their IPM needs case to be heard. It is curious that these same legislatures
that impose restrictions on pesticide use, and that consider the property posting and
other constraints on the landscape applicator (Anonymous 1988), don't ever seem to
include an IPM funding rider in their legislation to accommodate the new ways business
will have to be done in the landscape.
I estimate that the number of professional research and extension workers in orna-
mentals entomology, plant pathology, and related disciplines has approximately doubled
in the past ten years. In some cases new positions have been created, but in most
instances these new workers are established persons in traditional agricultural or forest
plant protection disciplines who for one reason or another have seen an opportunity in
ornamentals work. Yet even this new influx of talent is out of proportion to the mag-
nitude of the job to be done in ornamentals.
A third impediment to field implementation of ornamentals IPM is the shortage of
trained personnel skilled in innovative ways of putting IPM into practice (Davidson et
al. 1988). Ten to 15 years ago many institutions developed pest management curricula
aimed at producing BS or MS level IPM practitioners. Many of these programs are now
floundering, the result of a nationwide shortage of good students interested in a practical
career in field agriculture and horticulture. The new opportunities in biotechnology
seem much more exciting and attractive to good students.

394 Florida Entomologist 72(3) September, 1989

Despite all these impediments, one has to balance them against the strong, positive
forces driving IPM forward. Pest management in ornamentals can never return to its
state of 20 years ago, or even 10. It is extremely unlikely that individual components
of IPM programs, such as the use of pest-resistant ornamentals, or biological control,
will by themselves bring solution to a major number of ornamentals pest problems. The
only reasonable course into the future seems to be IPM, and I am completely confident
that this concept will continue to capture an increasing portion of the ornamentals pest


Anonymous. 1988. Sign-posting, licensing laws in the books. Landscape Mgt. 27(7): 8.
California Nature Conservancy. 1987. Sliding toward extinction: the state of Califor-
nia's natural heritage, 1987. Jones and Stoker Associates, Sacramento, Califor-
nia. 105 p.
DAVIDSON, J. A., C. F. CORNELL, AND D. C. ALBAN. 1988. The untapped alterna-
tive. Amer. Nurseryman 167(11): 99-109.
PARRELLA, M. P., AND V. P. JONES. 1987. Development of integrated pest manage-
ment strategies in floricultural crops. Bull. Entomol. Soc. Amer. 33: 28-34.
SMITH, D., AND S. GILL. 1987. Anatomy of an IPM program. Landscape Mgt. 26(8):
SMITH, D. C., AND M. J. RAUPP. 1985. Economic and environmental assessment of
an integrated pest management program for community owned landscape plants.
J. Econ. Entomol. 79: 162-165.


'Department of Entomology,
University of California, Riverside, California 92521
2Department of Biology,
Utah State University, Logan, Utah 84321
'Department of Horticulture,
Agriculture University, Wageningen, The Netherlands


The traditional viewpoint that there is a zero tolerance level for pests or their
damage on ornamental plants has hindered the development of statistically accurate
sampling plans. Monitoring procedures determining initial presence, peak flight times,
etc. have been developed, but actual methods to estimate pest populations on plants
are lacking. This is changing coincidentally with grower perceptions that some pests or
damage can be tolerated. This attitude change is most common in crops where a consid-
erable portion of early growth is not marketed, where pesticide resistance renders even
weekly insecticide applications less than satisfactory, or where restrictions on pesticide
use has forced growers to be more judicious about pesticide applications. Recent devel-
opments in sampling for adults and larvae of the leafminer, Liriomyza trifolii (Burgess)
(Diptera: Agromyzidae) using yellow traps and/or leaf samples in chrysanthemums,

Parrella et al.: Symposium-IPM for Ornamentals

marigolds and gypsophila are reviewed. In addition, the utility of using yellow traps to
monitor adult whitelifes, Trialeurodes vaporariorum (Westwood) and Bemisia tabaci
(Gennadius) (Homoptera; Aleyrodidae) on poinsettias, is discussed.


El punto de vista traditional de que hay un nivel de cero tolerancia hacia plagas o a
su dafio a plants ornamentales, ha sido un obstdculo en el desarrollo de muestreos
estadisticamente precisos. Se ha desarrollado el procedimiento para determinar el
chequeo de la presencia inicial, el auge del vuelo, etc., pero actualmente faltan m6todos
para estimar la poblaci6n de la plaga. Esto estA cambiando al mismo tiempo con la
percepci6n del productor de que algunas plagas o su daflo se pueden tolerar. Este
cambio de actitud es mAs comin en cultivos donde una porci6n considerable del primer
crecimientono es comerciable, cuando la resistencia a pesticides rinden ain aplicaciones
semanales de insecticides menos que satisfactorias, o cuando restricciones en el uso de
pesticides ha obligado al agricultor a ser mAs juicioso en la aplicaci6n de insecticides.
Se revisan recientes avances en el muestreo de adults y de larvas del minador,
Liriomyza trifoli (Burgess) (Diptera: Agromyzida), usando trampas amarillas y/o mues-
tras de hojas de crisantemos, clavelones y "gypsohila". Adicionalmente, se discute la
utilidad de usar trampas amarillas para chequear los adults de la mosca blanca,
Trialeurodes vaporariorum (Westwood) y de Bemisia tabaci (Gennadius) (Hom6ptera:
Aleyrodida) en poinsetias.

In California in 1986, the production environmental horticulture industry was valued
at greater than 1.3 billion dollars (Anon. 1988). Despite the size of this industry in
California and in the United States, limited information is available on IPM or sampling.
This is gradually changing with considerable research focusing on spider mites at-
tacking roses and foliage plants, whiteflies on poinsettia and leafminers attacking vari-
ous cut flowers and seed crops. One reason for greater concentration by researchers on
the development of sampling strategies is the realization (by both growers and resear-
chers) that pest damage on an ornamental crop can be tolerated without loss of aesthetic
(i.e., market) value. Prior to acceptance of this general concept, sampling plans were not
necessary; it was pointless to estimate pest densities when the only acceptable level
was zero or near to zero. Not all production ornamental crops can tolerate pest popula-
tions; however, there are specific situations when pests or their damage can be tolerated
by growers. These include situations where a considerable portion of early growth is
not marketed (e.g., many cut flower and seed crops). Furthermore, the decreasing
availability of efficacious insecticides together with legislative restrictions governing
their application will force growers to be more judicious about pesticide application.
Estimates of pest populations would permit growers to time their sprays with increasing
pest populations. This strategy may lead to an increase in the effectiveness of an insec-
ticide as well as a decrease in the probability of the development of insecticide resist-
ance. The alarming rate at which serious pests have spread worldwide throughout the
ornamentals industry [e.g., Liriomyza trifolii (Burgess), Spodoptera exigua Hiibner,
Frankliniella occidentalis (Pergande), and Bemisia tabaci (Gennadius)] emphasizes the
point that current prophylactic applications of pesticides are unsuccessful for controlling
Although the development of more sampling strategies in production ornamentals
is currently underway, this segment of the environmental horticulture industry is far
behind landscape trees and shrubs. In this division of the environmental horticulture
industry, monitoring pest populations to establish optimum treatment times has been
suggested for many years (Pritchard & Beer 1950, Koehler et al. 1965, Raupp, 1985


Florida Entomologist 72(3)

and references therein) and considerable work has been done relating pest population
levels to acceptable plant injury (Raupp et al. 1988). Monitoring and sampling in land-
scape trees and shrubs will not be considered in this paper; rather, emphasis will be
placed on recent developments in sampling in floriculture. In particular, studies with
larvae and adults of the leafminer L. trfolii on gypsophila (Gypsophila paniculata),
chrysanthemums (Dendranthema grandiflora) and marigolds (Tagetes popular) and
adult whiteflies (Trialeurodes vaporariorum (Westwood) and B. tabaci on poinsettia
(Euphorbia pulcherrima) will be discussed.

Liriomyza trifolii in Chrysanthemums

The first step in developing a sampling plan for larvae of this pest in chrysan-
themums grown as cut flowers was to establish a relationship between mined leaves
and aesthetic injury. This varied from grower to grower as well as during time of the
year. Each grower has his/her own idea as to what level of damage is acceptable in a
given situation. We found that this would vary with the historical quality of the crop
and problems associated with insecticide resistance in L. trifolii. Growers known for
producing perfect flowers, for example, rarely would settle for any damage. The resist-
ance level of this leafminer has been shown to vary considerably from location to location
(Haynes et al. 1986). The more difficult the leafminer is to control, the greater the crop
damage at harvest. In addition, where the flowers are sold also influences the level of
damage that is acceptable. Growers selling to a broker tolerate less damage than those
selling at local flower markets. Finally, acceptable damage fluctuates with the time of
year. During the holidays cut flowers are worth far more than at other times. Intui-
tively. it would seem that these holiday crops, which bring greater return to the grower,
must have little damage; however, just the opposite is true. Because the demand for
flowers is so great around these holidays, there is usually a flower shortage and growers
can sell all of their production regardless of the amount of the damage. This would not
be the situation if the flowers themselves were damaged in any way. During the summer
months there is an overabundance of product and chrysanthemums with mined foliage
will sell considerably below the market price.
Given the vagaries just described in establishing aesthetic injury levels, we felt that
it would be best to develop a binomial sampling plan that permitted a very rapid assess-
ment of the population with a minimum amount of work and did not require specifying
a critical level of damage in advance. Each grower could therefore decide what damage
level was acceptable and proceed to use population estimates as a guide to prevent
additional damage (Parrella & Jones 1987).

Liriomyza trifolii Larvae in Chrysanthemum

Chrysanthemums can be harvested in as little as 12 weeks, but during this period
the crop starts as small rooted cuttings approximately 10 cm in height and is greater
than 100 cm tall at harvest. Studies on the distribution of oviposition and feeding by L.
trifolii was conducted in the greenhouse as described by Parrella et al. (1987) Briefly,
a greenhouse section (6 m wide x 30 m long) was planted with 3,000 chrysanthemum
(cv 'Manatee Iceberg') plants arranged in 4 beds. Plants were grown as a pinched crop
and normal grower practices were followed. A total of 50 plants were randomly selected
within this area and upon planting, immediately caged to exclude flies. Cages were
constructed from clear acetate (1 mm thick) which was rolled into a 20 cm diam cylinder.
The edges of the cage were sealed with duct tape and the bottom of the cage was driven
into the soil to prevent fly entry. The tops and two 12 cm diam air holes on the sides
were covered with fine mesh cloth (200 holes/2.5 cm2) to allow air movement but to


September, 1989

Parrella et al.: Symposium-IPM for Ornamentals

exclude flies. No applications of pesticides were made in the study area until 11 weeks
after planting. Population levels in the greenhouse during this time were high (>200
flies per yellow trap per week, Parrella et al. 1987).
Each week, three caged plants were randomly selected and their cages removed;
one week later, plants were uprooted. Their roots were then placed in moist paper
toweling and the plants were transported to the laboratory. The stems on each plant
were ranked from tallest to shortest and, after 4 days, the plant, stem and leaf number
(using the youngest leaf on each stem as leaf number 1) were recorded along with the
number of feeding punctures and larvae or mines present on each leaf. The time between
plant removal and counting permitted hatching of eggs laid just prior to removal. Only
the 3 largest stems from each plant were used in the analysis because the other stems
on the plant had markedly fewer leaves.
The mean number of mines per leaf was calculated for all 9 stems per week. These
values were then used to calculate the percent of the total mines for that week found
on each leaf. Oviposition pattern exhibited by L. trifolii through the growth of the crop





10 20

Leaf Node

Fig. 1. Percent or total mines caused by Liriomyza trifolii on different leaf nodes
of chrysanthemum (leaf 1 is the youngest) during week 7-9. (N = 9 stems per week)


Florida Entomologist 72(3)

September, 1989

was consistent; they tended to avoid very old and very young leaves (Fig. 1). Con-
sequently, leaf samples taken from the middle portion of the plant are most likely to
contain larvae and provide the most reliable (and highest) estimate of leafminer popula-
tion density.
The development of sampling plans (utilizing leaf samples from the middle part of
the plant) was conducted in two greenhouses during the summers of 1982 and 1983.
Details are given in Jones & Parrella (1986a). An important aspect of the study was
that covariance analysis revealed that the coefficients of Taylor's power law were not
significantly different between plant strata or between crops with one or many cultivars.
Thus, a common regression could be used to describe the relationship between variance
and mean. A fixed sample size of 100 leaves provided a better estimate of mean popula-
tion densities than a constant precision sample (i.e., a sequential sampling design in
which sampling is terminated when a defined level of precision is achieved). A binomial
(presence-absence) sampling plan was developed to estimate leafminer densities rapidly
during crop growth as well as for total mining damage at harvest. The latter aspect
was completed so growers (or researchers) could rapidly assess the final damage to the
marketed portion of the chrysanthemum crop.

Liriomyza trifolii Larvae in Gypsophila

With the field grown flower crop, Gypsophila paniculata cv perfectt' (commonly
called gypsophila or baby's breath), the most prevalent leafminer attacking this crop in
California is the pea leafminer, L. huidobrensis Blanchard (Parrella & Bethke 1984).
Baby's breath is planted as rooted cuttings and generally flowers in 60 to 120 days,
depending on environmental conditions; peak flowering periods in California are from
February-October. The crop is routinely cut back and will reflower. Some growers
replant annually while others may keep plants as long as 5 to 6 years.
One coastal planting of gypsophila (70 x 160 m), consisting of ca. 6,000 plants, was
sampled for leafminers immediately after it was cut back until flowering. Twenty plants
were randomly chosen each week and 10 leaves were taken in a stratified design-10
from the bottom, vegetative part, 10 from the middle, bolting section, and 10 from the
upper, flowering section. All leaves were returned to the laboratory and examined for
feeding punctures and the presence of mines.
Data for punctures and mines plotted over the 14-week sample period clearly showed
a preference by L. huidobrensis for the lower, vegetative section of the gypsophila plant
(Figs. 2 and 3). A marketed bunch of baby's breath consists of the upper, flowering
section where there are few mines or punctures. In addition, L. huidobrensis mines the
lower leaf surface (Parrella et al. 1985) and the mines are usually not visible from the
upper side of the leaf. Therefore, leafminer control on gypsophila is usually not neces-
sary, especially when considering the large parasite complex associated with leafminers
in this crop (Price & Stanley 1982).

Liriomyza trifolii Adults in Chrysanthemum

Mark-release-recapture studies in a range of greenhouse chrysanthemums (Jones &
Parrella 1986b) provided insight into the movement of L. trifolii within a greenhouse
as well as information on proper yellow trap placement to obtain maximum data on
leafminer population dynamics. This study demonstrated that traps should be spaced a
minimum of 26 m apart for gathering information on population trends and 47 m apart
(between plots) for evaluation of treatment effects within plots (e.g., pesticide evalua-
Such information was used to develop constant precision sampling plans for yellow


Parrella et al.: Symposium-IPM for Ornamentals 3!

Of12A1 .i. -

100 120 140 160 180
100 120 140 160 180


Fig. 2. Total leaf punctures per leaf on Gypsophila caused by Liriomyza huidobren-
sis (10 leaves per strata, 20 plants per sample date)

traps (Parrella & Jones 1985). Trapping at several locations over a three-year period
revealed that the variance/mean relationship of catches was consistent from year to
year, location to location, and trap type to trap type. A requirement is that traps must


1 .0-



100 120 140 160 180



Fig. 3. Total mines per leaf on Gypsophila caused by Liriomyza huidobrensis (10
leaves per strata, 20 plants per sample date)


400 Florida Entomologist 72(3) September, 1989

be positioned only over homogeneous blocks of chrysanthemums (i.e., chrysanthemums
planted less than 30 days apart). Standard formulae (Taylor's power law and Iwao's
patchiness regression) were fit to the data. Iwao's patchiness regression best described
the variance/mean relationship and stop lines were generated with a fixed level of
precision of 0.25. Validation of the sampling plant demonstrated that only 18% of the
traps throughout the season needed to be counted to achieve the fixed level of precision.

Trapping and Damage

Although adult L. trifolii captured on yellow cards can provide the grower with
important information regarding population fluctuations in the greenhouse, correlating
these data to crop damage (i.e., mines) would increase the yellow card's utility in deci-
sion making. A consistent relationship is difficult to determine, probably because trap
catches may vary with environmental conditions, normal greenhouse operations, cul-
tivar location, type of greenhouse, and the delay between oviposition and when mines
become visible. Some of these factors may affect the number of mines in the leaves
differently than they affect trap catches. In addition, the number and type of cultivars
(which differ in susceptibility to leafminers) in the greenhouse are of critical importance.
In 1986, a correlation was run between adult L. trifolii caught on yellow cards and
live larvae present in leaves. Data were collected from a 10,000 ft2 greenhouse contain-
ing marigolds (Tagetes papula) grown for seed. There were 6 cards (7.5 x 12.5 cm) in
this greenhouse which were checked twice weekly. At the same time 18 plants were
randomly selected and one leaf was removed every 2.5-7.5 cm of plant height. Live
larvae were counted in these leaves immediately. Sampling was done over a 4-mo. time
period, from crop inception to harvest. Methods were similar in 1987 except that the
greenhouse was 6,700 ft2 and there were 12 plants per sample. A strong correlation was
found when leafminer populations were high (Fig. 4) (4 = 0.8556, P <0.001). However,
when populations were lower the following year in this same greenhouse (Fig. 5) there
was no correlation. Unfortunately, it is more important from a decision-making
standpoint to be able to use the yellow cards in a predictive sense when populations are

u.. F = 40.967, P << .01
S 10
0r = 0.8556

< 1- 6I

0 50 100 150 200 250 300 350 400 450 500


Fig. 4. Predictability of larval populations of Liriomyza trifolii in marigolds for
adults caught on yellow cards. Correlation calculated with live larvae abscissaa) time
lagged 7 days. This greenhouse represents high populations of adult leafminers.

1 .0





Parrella et al.: Symposium-IPM for Ornamentals 40

F = 3.665, P = .0725
r = 0.4213

I I I I_________

0 5 10 15 20 25

Fig. 5. Predictability of larval populations of Liriomyza trifolii in marigolds for
adults caught on yellow cards. Correlation calculated with live larvae abscissaa) time
lagged 7 days. This greenhouse represents low populations of adult leafminers.

low, before serious mining injury occurs. Based on our studies, yellow cards may not
function effectively in predicting damage when populations are low.

Whiteflies and Poinsettia

During the fall of 1987, research was conducted in a Christmas crop of poinsettias,
using yellow cards (hung 30 m over the crop) to trap both T. vaporariorum and B.
tabaci. These cards were checked weekly through the crop although only 3 data sets (3
dates) are presented here. There were 275 yellow cards hung uniformly in an area
encompassing ca. 0.4 ha.
A simple formula for calculating sample size which utilizes the standard error of the
mean as a measure of precision (i.e.. SE/k) (Southwood 1982, Karandinos 1976) was
applied to the trap catch data. We found the number of traps required to reach a
precision level of 0.25 was well below the number currently in use (Table 1) particularly
when the R whiteflies per card exceeded 4.0. Therefore, the grower could reduce the
number of traps and still be confident of population trends in the greenhouse. As the



Date Xa S.D.a E = 0.15 E=0.20 E = 0.25

10/01 10.5 12.2 228 130 84
09/14 4.0 8.0 683 384 245
10/01 2.0 3.8 614 345 222

"Estimated from 275 cards spaced uniformly over a 0.4 ha. poinsettia range. All cards checked weekly.
bEstimated from Southwood (1982) and Karandinos (1976) using: N = Z' 2) (S/EX)2 where N = samples required,
7Z/2 = 1.96 S = standard deviation, X = mean, E = predetermined standard error as a decimal of the mean.

Florida Entomologist 72(3)

number of traps in use declines, however the potential to detect 'hot' spots in the
greenhouse is also reduced. The grower must then make the decision as to the number
of traps to employ and any known hot spots should be trapped separately.
As yellow traps become more and more popular, there are some who advocate their
use in trapping out populations of whiteflies. This has been done in the Mediterranean
area in tomato greenhouses where yellow cards were utilized together with parasite
releases (van de Vrie & Vacante 1984). However, there have been no data published to
date on the utility of mass-trapping whitefly populations in ornamental greenhouses.


Monitoring insect populations in the greenhouse can provide a grower with the
following information:
1. Population trends in the greenhouse. Consequently, potential outbreaks can be
2. Whether populations build up in the greenhouse or migrate in from outside. If
migration is the main problem, screening can be considered.
3. The optimal time to apply pesticides. Sprays can be aimed at population peaks of
the most susceptible stages.
4. An evaluation of spray efficacy. If populations targeted by the pesticide drop after
application, then obviously the insecticide has been successful.
5. More intelligent, judicious use of pesticides based on population trends rather
than on a calendar schedule. This will result in less residue, environmental contamina-
tion and worker exposure. This is becoming increasingly important as rules and regula-
tions governing pesticides increase
6. A reduction in the probability of insecticide resistance developing in the target
pest through use of fewer pesticide sprays.
7. An overall monetary savings through reduction in pesticide use while maintaining
plant quality.


This research has been supported, in part, by the American Floral Endowment and
the California Association of Nurserymen.


ANONYMOUS. 1988. 1986 value of California nursery stock climbs above the billion
dollar mark. Pacific Coast Nurseryman 47: 16-18.
1986. Monitoring insecticide resistance with yellow sticky cards. California
Agric. 40: 11-12.
JONES, V. P., AND M. P. PARRELLA. 1986a. Development of sampling strategies for
larvae of Liriomyza trifolii (Diptera; Agromyzidae) in chrysanthemums. Envi-
ron. Entomol. 15: 268-273.
1986b. The movement and dispersal of Liriomyza trifolii (Diptera; Ag-
romyzidae) in a chrysanthemum greenhouse. Ann. Appl. Biol. 109: 33-39.
KARANDINOS, M. G. 1976. Optimum sample size and comments on some published
formulae. Bull. Entomol. Soc. Am. 22: 417-421.
KOEHLER, C. S., M. E. KALTOULAS, AND R. L. CAMPBELL. 1965. Timing of treat-
ments for control of irregular pine scale. J. Econom. Entomol. 58: 1102-110.5
PARRELLA, M. P., AND J. A. BETHKE. 1984. Biological studies of Liriomyza huidob-
rensis (Diptera: Agromyzidae) on chrysanthemum, aster and pea. J. Econ. En-
tomol. 77: 342-345.


September, 1989

Raupp et al.: Symposium-IPM for Ornamentals

PARRELLA, M. P., AND V. P. JONES. 1985. Yellow traps as monitoring tools for
Liriomyza trifolii (Diptera; Agromyzidae) in chrysanthemum greenhouses. J.
Econ. Entomol. 78: 53-56.
1987. Development of integrated pest management strategies in floriculture
crops. Bull. Entomol. Soc. Am. 33: 28-34.
PARRELLA, M. P., V. P. JONES, AND G. D. CHRISTIE. 1987. Feasibility of using
parasites for the biological control of Liriomyza trifolii (Diptera: Agromyzidae)
on commercially grown chrysanthemums. Environ. Entomol. 16: 832-837.
PARRELLA, M. P., V. P. JONES, AND L. M. LEBECK. 1985. Effect of leafmining and
leaf stippling of Liriomyza spp. on photosynthetic rates of chrysanthemums.
Ann. Entomol. Soc. Am. 78: 90-93.
PRICE, J. F., AND C. D. STANLEY. 1982. Gypsophila, leafminer and parasitoid re-
lationships on farms of differing pesticide use patterns, pp. 66-71, in Proc. 3rd
Annual Industry Conf. on the Laefminer, San Diego, Calif. Soc. Am. Florists.
Alexandria, VA. 216 pp.
PRITCHARD, A. E., AND R. E. BEER. 1950. Biology and control of Asterolecanium
scales on oaks in California. J. Econ. Entomol. 43: 494-497.
RAUPP, M. J. 1985. Monitoring: An essential factor to managing pests of landscape
trees and shrubs. J. Arboriculture 11: 349-355.
DORFER. 1988. Decision-making considerations for aesthetic damage caused by
pests. Bull. Entomol. Soc. Am. 34: 27-32.
SOUTHWOOD, T. R. E. 1982. Ecological methods. John Wiley & Sons, New York.
524 pp.
VAN DE VRIE, M. AND V. VACANTE. 1984. Greenhouse whitefly control through the
combined use of the color attraction system with the parasitic wasp Encarsia
formosa (Hym.: Aphelinidae). Entomophaga. 29: 303-310.



'Department of Entomology,
University of Maryland, College Park, MD 20742
2Cooperative Extension Service,
University of California, Berkeley, CA 94720
3National Resource and Environment Division
Economic Research Service, USDA, Washington, D.C. 20005


This article reviews decision-making considerations that apply to the management
of insect pests of ornamental plants. In particular we estimate a modified Economic
Injury Level (EIL) for the bagworm, Thyridopteryx ephemeraeformis (Haworth), at-
tacking American arborvitae, Thuja occidentalis under retail nursery conditions. Under
these circumstances, the EIL was found to be only about four first instar larvae per 4
ft tree. This confirms earlier suppositions concerning the low tolerance for pests causing
aesthetic injury. In addition, we briefly examine the concepts of Aesthetic Injury Levels

404 Florida Entomologist 72(3) September, 1989

(AIL) and Aesthetic Thresholds and illustrate how these may be estimated for pests
causing aesthetic injury.


En este articulo se revisan los factors utilizados en el process de toma de decisions
para el manejo de insects consideralos como plagas de plants ornamentales. En particu-
lar, hemos estimado un nivel de dafo econ6mico para el gusano cargacartucho Thyridop-
teryx ephemeraefozmis (Haworth), el cual ataca a Thuja occidentalis en planteles de
plants ornamentales. Bajo estas circunstancias, se determine un nivel de daio
econ6mico de alrededor de 4 larvas en el primer estadio por cada 4 pies de Arbol. Con
esto, se confirman suposiciones anteriores relacionadas a la baja tolerancia de plagas
causantes de dafios est6ticos. En adici6n, se examine brevemente el concept de niveles
de daio estetico y umbrales est6ticos y se ilustra como 6stos pueden ser estimados en
plagas que causan dafos est6ticos.

The last decade has seen a proliferation of integrated pest management programs
in residential environments (Olkowski et al. 1976, 1978, Davidson et al. 1981, Hellman
et al. 1982, Short et al. 1982, Holmes & Davidson 1984, Raupp & Noland 1984, Smith
& Raupp 1986, Cornell & Davidson 1987). One important limitation to several of these
programs was the lack of rigorously defined decision-making rules similar to the
economic injury level that has grained wide use in agronomic systems. Potter (1986)
and Pedigo et al. (1986) have identified the lack of decision-making guidelines as a
fundamental limitation of urban pest management programs.
One decision-maing approach that has gained widespread utility in agronomic sectors
is the use of economic injury levels (EIL). Stern et al. (1959) were the first to introduce
this term. They defined it as the lowest population level that causes economic damage.
Recently, Pedigo et al. (1986) presented a standardized model useful in determining
economic injury levels. Using this approach the EIL is determined by the cost of man-
agement per unit of production (C), the market value per unit of produce (V), the injury
units per insect for each unit of produce (I), and the damage per unit of injury (D). The
EIL is expressed in injury equivalents per unit of production and is related to these
parameters in the following way: EIL = C/VID.
This article serves a synopsis of a more thorough review published elsewhere (Raupp
et al. 1988). Detailed descriptions of methods, results, and their interpretation are
found in this earlier work. Here we briefly examine decision-making considerations as
they relate to pests of woody ornamental plants. To demonstrate how decision-making
rules can be formulated for pests causing aesthetic damage, we selected a common
ornamental plant American arborvitae (Thuja occidentalis) and an associated insect
pest the bagworm (Thyridopteryx ephemeraeformis (Haworth).
We used the model of Pedigo et al. (1986) to estimate an EIL for a defoliator, the
bagworm, on an ornamental shrub, American arborvitae. To determine the parameters
used to calculate the EIL, we adopted the perspective of a retail nurseryman who
manages ornamental plants for economic gain.
Using the estimates described in Raupp et al. (1988) and the methodology of Pedigo
et al. (1986), the EIL for this system was about 4 bagworm larvae/4 ft tree. From an
operational standpoint, this extremely low EIL has several important implications.
Published accounts of bagworm life histories in conjunction with our findings indicate
that a single female bagworm has the potential to create an infestation exceeding the
EIL of the retail nurseryman (Barrows 1974, Horn & Sheppard 1979). This finding
highlights the need for effective monitoring approaches that can facilitate pest detection

Raupp et al.: Symposium--IPM for Ornamentals

and intervention prior to injury. Furthermore, this finding confirms the supposition
made by several authors regarding the extremely low tolerance of people for aesthetic
injury (National Research Council 1980, Larew et al. 1984, Zungoli & Robinson 1984,
Sadof & Raupp 1987, Raupp et al. 1988).
The plant retailer is but one decision-maker with regard to managing pests of orna-
mental plants. The vast majority of woody ornamental plants are components of man-
aged landscapes. For pest managers such as homeowners, economic factors may not be
the primary ones that affect management decisions (Potter 1986, Pedigo et al. 1986,
Raupp et al. 1988). It was for these non-economic situations that Olkowski (1974)
suggested the use of an Aesthetic Injury Level (AIL) to form the basis for management
decisions. The AIL was considered to be the lowest level of a pest that caused aesthetic
injury (Olkowski 1974). This concept proved extremely valuable for making manage-
ment decisions for a variety of insect pests attacking street trees in California (Olkowski
et al. 1978).
Since the AIL was first conceived, several other works have contributed to our
understanding of decision-making as it relates to pests causing aesthetic injury. Buhyoff
& Leuschner (1978) provided one of the first examples of the relationship between plant
injury and aesthetic damage. They photographed forested landscapes that had been
injured by the southern pine beetle. These photographs were shown to different groups
of subjects that evaluated their landscape utility. The visual preference of those sur-
veyed declined dramatically between injury levels of 0 and 10% of the landscape dam-
aged. At greater levels of injury, little decline in disutility was noted. Based on their
findings, Buhyoff & Leuschner (1978) suggested that management efforts for southern
pine beetle be directed at preventing new outbreaks.
Koehler & Moore (1983) were the first to quantify the relationship between insect
abundance and an indicator of aesthetic quality. For the cypress tip moth, Argyresthia
cupressella Walsingham, they demonstrated that the unsightliness of a plant increased
as a simple linear function of tipminer abundance. Larew et al. (1984) determined a
similar relationship for serpentine leafminer injury on chrysanthemums. In this case
leafminer injury was directly related to the marketability of the plant.
Parrella & Jones (1987) also emphasized the importance of aesthetic considerations
in managing pests of greenhouse crops. Cut flowers, such as chrysanthemums and
gerbera, sustain considerable injury from leafminers. However, the injury often occurs
on plant parts that are not marketed. Parrella & Jones (1987) suggested that a modified
AIL be adopted for the floricultural industry. This AIL would permit damage to non-
marketed parts thereby reducing unnecessary pesticide use. Reductions in the number
of of pesticide applications could greatly facilitate biological control in greenhouses.
The ideas formulated in these earlier works prove useful in generating decision-mak-
ing rules for the system involving arborvitae and bagworms. During the survey at retail
nurseries, we asked customers which plant or plants showed damage and which plant
or plants had enough damage to warrant control. The first question was used to estimate
the bagworm density that caused aesthetic damage. We regard this density to be analog-
ous to the EIL defined by Stern et al. (1959). However, one difference is noteworthy.
In the case of the EIL, the density of importance is the lowest one causing economic
injury. In the case of the AIL, we were interested in establishing a density that would
be perceived as damaging by the average individual. The level of injury perceived as
damaging by half of those surveyed corresponded to about 5% of the leaf area missing
or discolored. This estimate translates into about 6 first instar bagworm larvae per 4
ft tree. For this system the EIL for the retail nurseryman and AIL for the customer
are quite similar.
Stern et al. (1959) defined the economic threshold as the population density at which
control is determined to prevent a population from reaching an economic level. The


Florida Entomologist 72(3)

relationship between plant injury and the perceptions of retail nursery customers can
be used to identify a similar decision-making rule based on aesthetic considerations. By
asking customers to identify which plant or plants had enough damage to warrant
control, we determined that half would initiate control when about 6% of the leaf area
was missing or discolored. The bagworm density causing this level of injury is about
nine per 4 ft tree. This density is functionally analagous to the ET defined by Stern et
al. (1959) and can be considered an aesthetic threshold (AT) for this system.
In some cases the mere presence of an offending organism may be sufficient to cause
injury and initiate a management decision. Wood et al. (1981) and Zungoli & Robinson
(1984) surveyed occupants of public housing units in three mid-Atlantic cities. Their
data indicate that as few as two cockroaches observed in an afternoon were perceived
as creating a problem by half of those surveyed. Furthermore, one to two cockroaches
seen in a household in a day were sufficient to warrant a control action by half of those
surveyed. These residents had well-defined perceptions concerning when coackroaches
were creating a problem and when they required control.
In summary, it is our hope that this review will prove useful in formulating decision-
making rules for pests causing aesthetic injury. Perceptions of those influencing man-
agement decisions must be quantified when the utility of the managed resource is based
on its appearance. When economic gain is important, as is the case of nursery and
greenhouse crops, economic injury levels and thresholds will form the basis of manage-
ment decisions. However, in many systems such as those involving established land-
scapes and turf, aesthetic consideration may overshadow economic ones. Aesthetic in-
jury levels and thresholds may be the appropriate basis for management decisions in
these systems.


We thank Ms. Elaine Mesavage who helped in preparing this manuscript. Computer
support was provided by the Computer Science Center at the University of Maryland.
We thank two reviewers for their helpful comments. This is Scientific Article No.
A-4769, Contribution No. 7773 of the Maryland Agricultural Experiment Station.


BARROWS, E. M. 1974. Some factors affecting population size of Thyridopteryx
ephemeraeformis (Lepidoptera: Psychidae). Environ. Entomol. 3: 329-332.
BUHYOFF, G. J., AND W. A. LEUSCHNER, 1978. Estimating psychological disutility
from damaged forest stands. Forest Sci. 24: 424-432.
CORNELL, C. F., AND J. A. DAVIDSON. 1987. The potential value of IPM to the
nursery industry: results of the Maryland pilot nursery IPM program. Part V.
M.S. Scholarly Paper. University of Maryland, College Park.
DAVIDSON, J., J. L. HELLMAN, AND J. HOLMES. 1981. Urban ornamentals and turf
IPM, pp. 68-72, in Proceedings of integrated pest management workshop. The
National Cooperative Extension, Dallas.
HELLMAN, J. L., J. DAVIDSON, AND J. HOLMES. 1982. Urban integrated pest man-
agement in Maryland, pp. 31-38, in H. D. Niemcyzk & B. G. Joyner [eds.],
Advances in turfgrass entomology. Hammer Graphics, Picqua, Ohio.
HOLMES, J. J., AND J. A. DAVIDSON. 1984. Integrated pest management for ar-
borists: implementation of a pilot program in Maryland. J. Arboric. 10: 65-70.
HORN, D. J., AND R. F. SHEPPARD. 1979. Sex ratio, pupal parasitism, and predation
in two declining populations of the bagworm, Thyridopteryx ephemeraeformis
(Haworth) (Lepidoptera: Psychidae). Ecol. Entomol. 4: 259-265.
KOEHLER, C. S., AND W. S. MOORE. 1983. Resistance of several members of the
Cupressaceae to the cypress tip miner, Argyresthia cupressella. J. Environ.
Hort. 1: 87-88.

September, 1989

Raupp et al.: Symposium-IPM for Ornamentals

LAREW, H. G., J. J. KNODEL-MONTZ, AND S. L. POE. 1984. Leaf miner damage.
Greenhouse Manager 3: 53-55.
National Research Council. 1980. Urban pest management. National Academy Press,
Washington, D.C.
OLKOWSKI, W. 1974. A model ecosystem management program. Proc. Tall Timbers
Conf. Ecol. Anim. Control Habitat Manage. 5: 103-117.
management: a framework for urban pest control. BioScience. 26: 384-389.
L. LAMB, AND L. ORTHEL. 1978. Pest control strategies: urban integrated
pest management, pp. 215-234, in E. H. Smith & D. Pimentel [eds.], Pest control
strategies. Academic, New York.
PARRELLA, M. P., AND V. P. JONES. 1987. Development of integrated pest manage-
ment strategies in floricultural crops. Bull. Entomol. Soc. Am. 33: 28-34.
PEDIGO, L. P., S. H. HUTCHINS, AND L. G. HIGLEY. 1986. Economic injury levels
in theory and practice. Annu. Rev. Entomol. 31: 341-368.
POTTER, D. A. 1986. Urban landscape pest management, pp. 219-252, in G. W. Ben-
nett & J. M. Owens [eds.], Advances in urban pest management. Van Nostrand
Reinhold, New York.
RAUPP, M. J., AND R. M. NOLAND. 1984. Implementing landscape plant management
programs in residential and institutional settings. J. Arboric. 10: 161-169.
DERFER. 1988. Decision-making considerations for pests causing aesthetic dam-
age. Bull. Entomol. Soc. Am. 34: 27-32.
SADOF, C. S., AND M. J. RAUPP. 1987. Customer attitudes toward defoliation of
American Arborvitae, Thuja occidentalis, by bagworm, Thyridopteryx
ephemeraeformis. J. Environ. Hort. 5: 164-166.
SHORT, D. E., J. A. REINART, AND R. A. ATILANO. 1982. Integrated pest manage-
ment for urban turfgrass culture-Florida, pp. 25-30, in H. D. Niemczyk & B. G.
Joyner [eds.], Advances in turfgrass entomology. Hammer Graphics, Picqua,
SMITH, D. C., AND M. J. RAUPP. 1986. Economic and environmental assessment of
an integrated pest management program for community owned landscape plants.
J. Econ. Entomol. 79: 162-165.
integrated control concept. Hilgardia 29: 81-101.
WOOD, F. E., W. H. ROBINSON, S. K. KRAFT, AND P. A. ZUNGOLI. 1981. Survey
of attitudes and knowledge of public housing residents towards cockroaches.
Bull. Entomol. Soc. Am. 27: 9-13.
ZUNGOLI, P. A., AND W. H. ROBINSON. 1984. Feasibility of establishing an aesthetic
injury level for German cockroach pest management programs. Environ. En-
tomol. 13: 1453-1458.


408 Florida Entomologist 72(3) September, 1989


University of Florida, IFAS
Central Florida Research and Education Center Apopka
2807 Binion Road, Apopka, FL 32703

University of Georgia
Agricultural Experiment Stations
Georgia Station, Griffin, GA 30223, U.S.A.


The potential for developing biological control programs for ornamental plants is
discussed. Biological control is a viable control tactic in certain production systems.
These systems include crops in which the marketed product is sold without the damaged
tissue and in stock beds from which propagative material is obtained. The factors that
are impeding the implementation of control programs are also discussed.


Se discute el potential de desarrollar un program de control biol6gico para plants
ornamentales. El control biol6gico es una tactica de control viable en ciertos sistemas
de producci6n. Estos sistemas incluyen cultivos en los cuales el product comerciable
es vendido sin el tejido dafado y en planteles de los cuales se obtiene el material de
propagaci6n. Se discuten los factors que impiden la implementaci6n de programs de

There are over 140 species of insects and mites that are known pests in greenhouses
(Pritchard 1949). In recent years, the major pests, including aphids, leafminers, thrips,
mealybugs, whiteflies, and spider mites have generally been controlled by insecticides
and acaricides. However, it is becoming increasingly clear that the strategy of unilateral
reliance on chemical control will not be the final solution to pest problems. In this
regard, there are four major problems attendant to chemical control: (1) development
of resistance to chemicals in target pest species; (2) the dwindling supply of useful,
registered insecticides and acaricides and increased governmental regulation; (3) the
damaging (or detrimental) effect of these chemicals on nontarget species resulting in
secondary pest outbreaks; and (4) phytotoxic reactions by treated plants. On the other
hand, unilateral reliance on biological control should not be viewed as a sound strategy
because biological control alone does not always give adequate protection.
A solution to this problem lies in the utilization of integrated control (Stern et al.
1959) or what is now called integrated pest management (IPM). This is a management
system in which ecologically suitable and economically sound control tactics are em-
ployed to maintain pest populations at tolerable levels. With respect to the current
situation in many commercial greenhouses, there is a critical need for integrating biolog-
ical control agents with existing cultural and chemical control methods.
Biological control programs are currently available for the control of twospotted
spider mite and greenhouse whitefly on certain greenhouse grown vegetable crops, but

Osborne & Oetting: Symposium-IPM for Ornamentals 409

these programs have not been transferred successfully to ornamental production sys-
tems. Secondly, many of the remaining pest species, particularly thrips, aphids, leafmin-
ers, mealybugs, scales, and various species of Lepidoptera still require considerable
research before biological control programs can be implemented.
Although many of these pests have effective natural enemy complexes outside the
greenhouse, the effectiveness of a given natural enemy under greenhouse conditions
may be limited or as is often the case they may even fail to become established. The
opposite situation can occur as well: natural enemies which are ineffective under field
conditions may be in the greenhouse. For example, the parasite Encarsia formosa
Gahan is effective against greenhouse whitefly, Trialeuroides vaporariorum
(Westwood), in the greenhouse; however, the same species is generally ineffective in
many field situations, especially where temperature extremes either favor the pest or
cause direct mortality of the parasites. The underlying reasons for the differential per-
formances of natural enemies have not been identified. However, certain greenhouse
conditions may be quite favorable for one species of natural enemy, but not for another.
In his 1988 review of biological pest control in greenhouses, van Lenteren (van
Lenteren & Woets 1988) presented a scheme for selecting natural enemies to be used
in greenhouse IPM programs. A list of criteria was presented for use in preintroductory
evaluations of natural enemies. The importance of the criteria differed depending on
the type of release program to be used: seasonal inoculative or inundative.
It is our view that, for effective biological control programs to be implemented
successfully in the greenhouse, a proper attitude on the part of the grower is necessary.
Colleagues in New York have also noted the importance of this aspect (see Tauber 1977,
Tauber & Helgesen 1978). Some philosophical commitment to biological control including
the integration of biological and chemical control is necessary. A certain amount of
patience and a firm resolve for making biological control work is needed. In our experi-
ence, one of the foremost obstacles to the implementation of biological control in the
greenhouse has been a negative attitude on the growers' part. Presently, we nave noted
a swing toward the opposite extreme. Many growers have unrealistic expectations for
this control technique. Researchers have found themselves in a situation where growers
are trying to implement programs that are far more ambitious than knowledge or cur-
rent technology can support and transferring technology from field situations or from
vegetable crops to ornamental production hasn't always proven successful. Although
we can't meet the current demand for biological methods to aid in ornamental pest
management programs, there is an exciting thrust by industry to support research
throughout the United States to meet the demand.
Examples of systems which are being targeted for our initial thrusts include crops
in which the marketed product is sold without the damaged tissue, such as marigolds
which are grown for seeds (Heinz et al. 1988) or cut flowers such as chrysanthemums
(Parrella et al. 1987a) and roses (Sabelis 1981). Another system which seems amenable
to biological control is stock beds where cuttings are either sold to other producers or
used within the operation. It is in these systems that resistant populations are most
likely to develop because of the multiple pesticide applications. If a resistant pest de-
velops in an operation that sells cuttings, the chances of distributing this resistant strain
to other growers is great, thus endangering the entire industry.
In the ornamental industry there is the perception that plants must be grown with-
out any pests present, and that plants become unmarketable if pests are present or if
feeding damage is visible. It is a realistic goal to obtain clean plants if pesticides are
effective and being applied properly. However, the chemical use needed to maintain
this zero threshold is highly conducive to the development of pesticide resistant pest
populations in greenhouses (Parrella et al. 1987b). The problem we are confronting is

410 Florida Entomologist 72(3) September, 1989

that these extreme and exacting standards are not being satisfied with pesticides today.
A few visits to retail outlets will confirm that these criteria are not being met. Consum-
ers are inheriting the growers failures. Plants are being shipped that infested with
resistant pests and consumers are not prepared to handle these formidable problems.
The pests we can't control are often not controlled by the types of pesticides available
to the consumer. Most of the chemicals which enable a grower to produce quality plants
are restricted or not readily available to anyone but commercial producers.
Another aspect of this problem is that we distribute our failures worldwide. If a pest
of ornamentals develops resistance, it gets shipped. This has occurred with many differ-
ent pests including mites, aphids, thrips, leafminers, and whiteflies. The most dramatic
case has occurred recently with the sweetpotato whitefly, Bemisia tabaci. This insect
was first reported in Florida in 1894 but wasn't considered a pest until the fall of 1986.
During the fall of 1986, ornamental growers throughout Florida experienced major
losses as the result of B. tabaci. At that time, ornamental growers in the rest of the
world and Florida's vegetable industry hadn't reported any problems with this insect.
By late 1988, this "strain" of the sweetpotato whitefly appears to have found its way
into greenhouses throughout most of the world and Florida's vegetable growers have
blamed this pest for losses in excess of 20 million dollars.
The ultimate goal of the ornamental industry is to sell products of the highest quality
and to do this without the constant disruptions we have seen in the past few years due
to the development of resistant pests or the removal of effective pesticides from the
market. The programs we are proposing are aimed at using biological controls in a
portion of the production system where some damage can be tolerated and reserve the
use of pesticides for the final stages of the production process. Theoretically, these
tactics will enable us to reduce the pesticide pressure exerted on a "captive" pest
population and delay the development of resistance to pesticides, thereby preserving
one of the ornamental industry's most valuable resources-labeled pesticides.
Research is needed to document the level of control currently obtainable in the
production of ornamental plants with pesticides so that we have a standard against
which we can compare the control achieved with integrated programs. Secondly, larger
scale studies are needed. Research has demonstrated that pests such as mites and
whiteflies can be controlled but these studies have been conducted in research
greenhouses using a limited number of plants. These results may or may not be directly
applicable to commercial production houses.
Another critical need in the ornamental industry is the development of monitoring
programs that will enable growers to detect the presence of pests before pest popula-
tions reach levels causing economic damage. Most growers don't practice any form of
monitoring. These programs must be implemented by growers using all types of pest
management systems. The system can be as sophisticated as those being developed by
Jones and Parrella (1986), or they can be simple and tailored to the specific needs and
resources of the grower.
There are a number of other factors that limit the implementation of biological
control programs for ornamental plants. Because of space limitations we will briefly
discuss only the following factors: 1) limited availability of beneficial, 2) specific use of
recommendations, 3) multicropping (mixed crops), and 4) funding.

1) Limited commercial availability of effective beneficial:
This aspect covers many of the major impediments to implementing biological control
programs. First, there are only a few companies that produce beneficial insects and
mites. These companies are, with few exceptions, located in foreign countries or Califor-
nia. As a result, growers in the eastern United States who wish to purchase parasites
or predators must obtain them through the mail. Unfortunately, little is known about

Osborne & Getting: Symposium-IPM for Ornamentals 411

what effect shipping or stress might have on beneficial but it certainly reduces their
utility. Because the ornamental industry is so widespread throughout the United States,
insectaries that can supply regional needs or act as distributors of quality beneficial
for larger producers may be a possible remedy. This arrangement also would address
another critical need which is the lack of an advisory service that could help growers
implement or utilize this more sophisticated technology.
The lack of available beneficial also means that there is a limited selection. There
are no commercially available beneficial for a number of important pests and for most
major pests, there is only one beneficial insect or one beneficial mite available commer-
cially. It seems unrealistic to expect one organism to be effective under the many
diverse situations that occur in the ornamental industry. It has been established that
E. formosa has problems parasitizing whiteflies on certain varieties of cucumbers be-
cause the trichomes interfere with the searching process (Ponti et al. 1986). In addition,
the poor performance of this parasite during the winter months has resulted in a search
for parasites that are more effective at lower temperatures (Vet 1980).
Environmental constraints also have resulted in the search for additional predators
of T. urticae. Phytoseiulus persimilis A.H. is sensitive to high temperatures, medium
to low relative humidities and pesticides. Under these conditions the efficiency of this
predator is reduced and in some cases it has failed to become established. P. longipes
has been evaluated on a limited basis for the control of mites in interior plantings where
relative humidity prevents P. persimilis from reproducing. The initial results indicate
that P. longipes would be better suited for such environments.
The concept of releasing two or more different beneficial into the same system has
been debated. In an ornamental system where we are using the biological agents as
biological insecticides, we do not depend on them to become permanently established
in the greenhouse. Therefore, the arguments against multiple releases in classical
biological control programs do not apply. We are currently evaluating multiple releases
of four different species of predatory mites in commercial greenhouses. Multiple releases
increase our chances of having the beneficial become established and, in time, obtaining
control of T. urticae.

2) Specific use recommendations:
The present demand for information is greater than what is available. Commercial
growers are looking for specific recommendations and without step by step guidance
many are reluctant to incorporate biological control into their pest management pro-
grams. Because of budget and time constraints, researchers who are involved in de-
veloping biological control strategies normally are restricted to working with a single
pest on a single crop. Commercial growers are dealing with far more complex situations.
Currently, because of poor funding, cooperation and communication between resear-
chers, there isn't a single example of a successful IPM program.

3) Multiple Cropping:
Growing more than one plant type in a greenhouse or even on the same bench is a
common practice. Large monocultures, which are common in most of agriculture, are
the exception in the ornamental industry. In Florida, some growers produce over a
hundred different plant species and cultivars. This situation greatly complicates the
development of pest control programs regardless of which tactics are used to manage
the pest populations. Each species of plant has its own unique complement of pests and
its own sensitivities to environmental conditions, including phytotoxic reactions to pes-
ticides. Mixed cropping can prevent the development of a single comprehensive control
program that can be applied to an entire greenhouse: each crop must have its own
unique program.

412 Florida Entomologist 72(3) September, 1989

4) Funding:
The idea that research, if adequately funded, can provide solutions to all pest prob-
lems is commonly expounded. The level of funding for the development of biological
control programs certainly ranks as a major limiting factor but, as has already been
discussed, it is not the only one.
The lack of information mentioned earlier, can be explained in part by the lack of
funding devoted to developing alternative methods. The cost to develop one pesticide
into a commercial product can be as much as 30 million dollars. This amount of funding
has not been allocated to developing pest control of any form in ornamentals. Organiza-
tions that provide the funding needed to conduct the research and field trials normally
do not give research on ornamental pest problems a high priority.
In Florida, pest complexes of ornamentals are frequently pests of crops grown for
food and fiber, such as leafminers, aphids, thrips and whiteflies. If the use of pesticides
is not reduced, the development of pesticide resistant strains will continue in
greenhouses and will eventually move to other agronomic crops.
A strong case can be presented for developing microbial controls. Funding should
not be as limited as it has been for research on parasites and predators because patho-
gens that kill insects or mites can be commercialized and sold in a manner similar to
that used for chemical pesticides. For example, Bacillus thuringiensis is sold by a
number of companies. More recently, a product called Nolo bait is being marketed for
grasshopper control. The active ingredient is a micro-organism, Nosema locustae, that
kills grasshoppers and some crickets. Many companies are interested in developing
similar products, and it will not be long before many more reach the market place. In
general, the shelf life of these microbial pesticides will be shorter than conventional
pesticides but it will be longer than other types of biological control agents. Additionally,
the use of parasites, predators, and microbial pesticides will require more education
and training of the grower than has been provided for the use of conventional pesticides.
The availability of insect and mite pathogens may be no better than traditional
biological-control agents. The number of products will depend on the difficulty of regis-
tering microbial pesticides. EPA considers them as pesticides and therefore they are
subject to the same complicated registration process. Secondly, micro-organisms will
not be allowed entry in the U.S., if they are exotic, without appropriate permits.
Quality control will be somewhat easier to maintain because techniques are available to
quickly evaluate quality.
Although there are many obstacles, biological-control programs for ornamentals are
a viable technique that must receive additional funding and research so that they can
be integrated with other methods for managing pests of ornamentals.


This Florida Agricultural Experimentation Station Journal Series No. 9854.


DE PONTI, O. M. B., J. C. VAN LENTEREN, AND M. W. SABELIS. 1986. Breeding
cucumbers and tomatoes for improved biological control of whiteflies and spider
mites. HortScience 21: 603.
HEINZ, K. M., J. P. NEWMAN, AND M. P. PARRELLA. 1988. Biological control of
leafminers on greenhouse marigolds grown for seed. California Agriculture.
42(2): 10-12.
JONES, V. P., AND M. P. PARRELLA. 1986. The development of sampling strategies
for larvae of Liriomyza trifolii in chrysanthemums. Environ. Entomol. 15: 268-

Nielsen: Symposium--IPM for Ornamentals 413

LENTERN, J. C. VAN, P. M. J. RAMAKERS, AND J. WOETS. 1980. World situation
of biological control in greenhouses, with special attention to factors limiting
applications. Med. Fac. Landbouww. Rijksuniv. Gent. 45: 537-544.
LENTERN, J. C. VAN, AND J. WOETS. 1988. Biological and integrated pest control
in greenhouses. Ann. Rev. Entomol. 33: 239-269.
ALI. 1987a. Resistance to insecticides: What a grower can do. Grower Talks
50(11): 136-142.
PARRELLA, M. P., V. P. JONES, AND G. D. CHRISTIE. 1987b. Feasibility of parasites
for biological control of Liltiomyza trifolii (Diptera: Agromyzidae) on commer-
cially grown chrysanthemum. Environ. Entomol. 16: 832-837.
PRITCHARD, A. E. 1949. California greenhouse pests and their control. Calif. Agri.
Exp. Stn. Bull. 173: 1-72.
SABELIS, M. W. 1981. Biological control of twospotted spider mites using phytoseiid
predators. I. Agric. Res. Report 910. Pudoc, Wageningen, The Netherlands.
integrated control concept. Hilgardia 29: 81-101.
TAUBER, M. J. 1977. Problems and promise of biological control in protected culture
crops, pp. 95-96, in R. E. Webb and F. F. Smith [eds.], Pest Management in
Protected Culture Crops. Washington, D.C.: ARS-NE-85.
TAUBER, M. J., AND R. G. HELGESEN. 1978. Implementing biological control sys-
tems in commercial greenhouse crops. Bull. Entomol. Soc. Am. 24: 424-426.
VET, L. E. M. 1980. Laboratory studies on three Encarsia spp. and one Eretmocerus
sp. (Hym., Aphelinidae) parasites of the greenhouse whitefly Trialeurodes vap-
orariorum (Westw.) to access their efficiency as biological control. Med. Fac.
Landbouww. Rijksuniv. Gent. 45(3): 555-561.


Department of Entomology
1680 Madison Avenue
Wooster, OH 44691


Few native species of so-called pest organisms are capable of overwhelming vital
woody plants to the point of causing significant damage. There is a larger number of
secondary-action organisms that exploit weakened trees and shrubs, causing their
further decline and eventual death. Entomologist and phytopathologists might be more
helpful to the plant production and maintenance industries by adjusting their major
thoughts from insects and pathogens to tree adaptation and adaptibility. Then, instead
of considering how to directly control pest organisms we can begin to think in terms of
what can be done for trees to reduce their susceptibility and vulnerability to pests.
Classic tree improvement programs should be continued but require better coordina-
tion among disciplines for success. Traditionally, these programs have not received
significant funding from Federal or State governments. An immediate and productive
avenue for exploiting natural resistance might be to determine the influence of plant
vitality on insect/disease-plant relationships. This work must be done in cooperation
with plant physiologists to define physiological parameters associated with plant vital-

414 Florida Entomologist 72(3) September, 1989

ity. We must also work with landscape horticulturists to determine which cultural prac-
tices reduce vulnerability of landscape plants to important herbivores. A working
hypothesis for this effort could be that most organisms that threaten survival of land-
scape plants are opportunists that exploit weakened hosts. Although the validity of this
hypothesis requires rigorous testing, it is a conservative beginning, since there is evi-
dence that improving tree vitality reduces impact of insects and fungal pathogens.


Pocas species nativas de los tal llamados organismos plagas son capaces de abrumar
arboles hasta el punto de causar un dano significant. Hay un mayor nfmero de organis-
mos de acci6n secundaria que explotan Arboles y arbustos d6biles, causando decadencia
adicional y eventualmente la muerte. Entomologistas y fitopatologistas pudieran servir
de mayor ayuda a las industries productoras y de mantenimiento de plants, ajustando
sus ideas principles sobre insects y pat6genos, a su adaptaci6n y adaptabilidad. En-
tonces, en vez de considerar c6mo controlar directamente los organismos de plaga,
pudieramos empezar a pensar en thrminos de qu6 se puede hacer por los Arboles para
reducir su susceptibilidad y vulnerabilidad a las plagas.
Programs clasicos de mejoramiento de Arboles deben de continuar pero requieren
mejor coordinaci6n entire las disciplines para triunfar. Tradicionalmente, esos programs
no han recibido significantes fondos de los gobiernos Federales o Estatales. Un medio
inmediato y productive para explotar resistancias naturales pudiera ser el determinar
la influencia de la vitalidad de las plants en las relaciones entire el insecto y la plant
enferma. Este trabajo debe de ser hecho en cooperaci6n con fitofisidlogos para definir
los parametros fisiol6gicos asociados con la vitalidad de la plant. Tambien debemos
trabajar con horticulturistas para determinar las practices culturales que reduce la
vulnerabilidad de plants a herbivoros importantes. Una possible hip6tesis de este es-
fuerzo seria que la mayoria de los organismos que amenazan la sobrevivencia de plants
son oportunistas que explotan los hospederos debilitados. Aunque la validez de esta
hip6tesis require una prueba rigurosa, es un comienzo conservador, puesto que hay
evidencia que mejorando la vitalidad del abol se reduce el impact de insects y
pat6genos de hongos.

Most plant taxa are resistant to most pest organisms. Many landscape managers are
unaware of this fact or forget this truism as they strive to keep pests from reducing
the aesthetic quality or the vitality of urban plants.
Only a few native species of so-called pest organisms are capable of overwhelming
vital woody plants to the point of causing significant damage. There is a larger number
of secondary-action organisms that exploit weakened trees and shrubs, causing their
further decline and eventual death.
Entomologists and phytopathologists who work with the landscape and nursery in-
dustries sometimes become impressed with the ubiquity and severity of insect and
disease problems in production and maintenance of landscape plants. The perspective
of landscape horticulturists is often quite different. They are encouraged by the success
of a wide variety of plant materials over large geographical areas with various degrees
of environmental stress. Entomologists and phytopathologists might be more helpful to
the plant production and maintenance industries by adjusting their major thoughts from
insects and pathogens and their exploitive abilities to a consideration of trees and their
adaptation and adaptability. Then, instead of considering what can be done directly
against pest organisms we can begin to think about what can be done for trees to reduce
their susceptibility and vulnerability to pests. Much cooperative research is required to
understand insect tree-host relationships at a level that will permit and encourage plant

Nielsen: Symposium-IPM for Ornamentals

health care for landscape plants. This research thrust is only recently receiving critical
effort in Europe and North America.
Most discussions of host plant resistance deal with inherent susceptibility or exploita-
bility of a host by a pest. A workable definition of susceptibility for our symposium
might be the degree to which a vigorous plant species or cultivar (= cultivated variety)
can be exploited. This is a genetic trait that can be measured through comparative
studies of selected taxa. Our work with white-barked birches, Betula spp., and the
native bronze birch borer, Agrilus anxius Gory, is an example of this kind of study
(unpublished). All trees were subjected to identical conditions in one location and mon-
itored yearly for nine years. The native paper or canoe birch, Betula papyrifera Marsh.,
was relatively more resistant to this borer than any of the Asian or European birches
A follow-up study is underway in Michigan to evaluate the influence of environmental
stress on the physiological performance and vulnerability of paper birch to damage by
bronze birch borer. In this case, the experiment was designed to evaluate the impact
of stress treatments on the vulnerability of a relatively resistant taxon.
So, vulnerability is related to vitality and can be defined as a plant's likelihood to
be exploited and damaged by an herbivore at a given point-in-time. Vulnerability is a
manifestation of the interaction between a genotype and its surroundings.
Although the difference between susceptibility and vulnerability may seem some-
what trivial and inconsequential, this difference is the main theme for my remarks here.
Classically, we consider the potential for exploiting natural resistance through breeding,
evaluation, and selection of species and cultivars. And, the results are often unequivocal.
For example, our study of birches indicated that the native paper birch should be used
in North American landscapes whenever a white-barked birch is specified.


The classical approach to tree improvement has merit and deserves continued re-
search effort as conducted by F. S. Santamour, Jr., Director of the USDA's program
for genetic improvement of urban trees, at the U.S. National Arboretum in Washington,
D. C. However, this kind of work has not been popular and has not received significant
financial support at most research institutions.
The search for superior amenity trees probably began hundreds of years ago. The
cultivars London plane tree, Platanus acerifolia (Ait.) Willd., Lombardy poplar
Populus nigra 'Italica', and copper beach, Fagus sylvatica 'Atropunicea', were de-
veloped in the 17th century (Li 1963). Plant exploration and introduction became popular
during the 18th century. In the more recent past, most improvements in woody plants
have resulted from selection of superior types through provenance studies, primarily
with commercial forest species (Wright 1976). Little actual breeding has occurred with
shade trees; most of the effort with woody plants has been with shrubs, including roses
(Egolf 1968).
According to members of the Genetic Working Group of the Consortium for Environ-
mental Foresty Studies, efforts to identify and develop superior trees for landscape
usage has suffered because research efforts are "dispersed among many species and
many characteristics, with insufficient concentration to effect substantial improve-
ments," (Anonymous 1982). They recognize there is need to agree upon one or more
clear goals to allow efficiency of effort in urban tree improvement programs.
Most species and cultivars of woody plants used in urban environments are not
adapted to urban stresses, including air pollution, wind tunneling, radiant heat, soil
type and compaction, limited planting and growing space, deicing salts, and urine from

Florida Entomologist 72(3)

September, 1989

domestic animals. It is apparent that the greatest need for future urban forestry is
development of cultivars that can tolerate these and other urban stresses while main-
taining vitality. We need trees and shrubs with the inherent genetic capacity to survive
and remain vital with minimal maintenance.
This goal of genetic improvement of urban trees is appropriate for the 21st century
and beyond. Until this goal has been reached there is immediate need to begin providing
existing landscape plants with the inputs they need to optimize their vitality. Further-
more, information concerning which trees are best adapted to urban conditions, proper
site selection and tree planting methods, and relative resistance to important pest or-
ganisms needs to be assembled and packaged for distribution to user groups ranging
from landscape architects to practicing arborists. Regional research groups in tree im-
provement, landscape horticulture, phytopathology, and entomology could coordinate
this effort. Kielbaso & Kennedy (1983) suggested that the success of future urban pest
management will depend upon an effective, coordinated technology transfer program.
This effort must be interdisciplinary.


An immediate and productive avenue for exploiting natural resistance might be to
determine the influence of plant vitality on insect-plant relationships. To do this we
must first work with plant physiologists to define physiological parameters associated
with vitality and then work with landscape horticulturists to determine which cultural
practices reduce vulnerability to important herbivores.
A working hypothesis for this effort could be that most organisms that threaten
survival of landscape plants are opportunists that exploit weakened hosts. Stated
another way, enhancement and maintenance of tree and shrub vitality will reduce the
vulnerability of the urban forest to pest organisms. Although the validity of this
hypothesis requires rigorous testing, it is a conservative beginning, since there is some
evidence that improving tree vitality reduces the impact of insects and fungal patho-
Waring & Pitman (1983) demonstrated that thinning lodgepolep ine, Pinus contorta
Var. latifolia Engelm., stands increased the productivity of remaining trees and de-
creased their vulnerability to the mountain pine beetle, Dendroctonus ponderosae Hop-
kins. In this case, stagnated, overstocked stands were extremely vulnerable to the
beetle until competition was reduced through thinning. Results of this research have
generated a massive program in the Northwest to thin stagnated stands of lodgepole
and ponderosa pine, Pinus ponderosa Douglas, to reduce beetle kill. These efforts to
protect green belts along highways in Central Oregon are impressive. Other bark bee-
tles and some defoliators have shown preference for subvital hosts (Raffa 1986). There
are numerous examples in the phytopathology literature of stress-related disease syn-
dromes (Schoeneweiss 1986), including pine wilt disease, hardwood declines, and
perhaps Dutch elm disease.


If this "insects as opportunists" hypothesis has merit, how can it be used now to
reduce landscape pest problems and how can it be examined efficiently through the
research process to increase our understanding of insect-plant relationships?
Since the early 1980's some arborists have begun marketing integrated pest manage-
ment as part of tree health care programs. These programs are based on the premise
that enhancing tree vitality and controlling incipient pest infestations and infections are

Nielsen: Symposiumr-IPM for Ornamentals

more cost-effective and environmentally responsible than preventive spray programs.
While surveying and monitoring serve as the foundation of these efforts, targeted pes-
ticide applications and cultural tactics are used to maintain the vitality of key landscape
plants. Pruning, aerification, mulching, watering, and fertilization are becoming integral
parts of services provided by some arboricultural firms. The change from using cover
sprays for controlling pests to responsible landscape horticulture has been slow but will
accelerate, since liability insurance for pesticide applicators has increased dramatically.
Impending governmental decisions designed to reduce pesticide usage in the urban
environment may also hasten this transition.
Unfortunately, little research has been done to measure the impact of cultural tactics
on plant vitality and associated pest organisms. Carrow and Graham's (1968) work is
an exception. They found that the type of nitrogen fertilizer used determined the re-
spose of a sucking insect. We need more emphasis on this kind of work. Certainly the
time has come for this effort to begin.
Raffa (1986) pointed out that, "a control strategy must first provide consistent
economical plant protection to ever be adopted." The tree health care concept is being
used now by a small number of practitioners to reduce damage by insects and other
pests in existing urban and suburban forests while at the same time reducing reliance
on pesticides. This approach is proving to be efficient.
Natural resistance can be exploited further by increasing diversity of urban plan-
tings. Landscape horticulturists recommend some kind of formula like to 10-5 rule (Phil-
lips 1979) or the 20-10 limit (Bridgeman 1979) to increase diversity. These schemes
suggest that any one plant family should not compromise more than 10-20% of a city's
street population and that not more than 5-10% should be comprised of a single genus.
Diversity can also be increased by maintaining uneven aged stands within the urban
forest. This practice may not be tolerated within a city block or perhaps on a boulevard.
But, it could be implemented throughout a municipality or in an alternating design even
in smaller areas.

If we wish to improve understanding of insect-plant relationships and how we can
help plants help themselves against pests, we must begin by investigating the basic
biochemical processes of plants, how they change in response to urban stress, and how
herbivores respond to these changes.
In Raffa's (1986) summary for using plant defense in pest management programs,
he stated: "From a theoretical standpoint we need to better understand how the various
components of co-adapted plant-insect associations interact. We can best develop this
understanding by emphasizing the dynamic aspects of these relationships under natural
conditions, and by manipulating key processes under controlled conditions."
Once we begin to understand how cultural practices and herbivory alter the dynamic
process of vulnerability, we can begin maximizing exploitation of natural resistance as
a management tactic for landscape plants. Of course, the best landscape management
strategy, even then, will result from functional integration of landscape horticulture,
weed science, phytopathology, and entomology. In the meantime, we all need to make
an effort to extend results of our applied and basic research programs to user groups.
We also need to develop closer ties to the production and landscape industries to have
more impact on what is produced and how landscape management is conducted. These
linkages might be difficult to achieve as a new generation of university researchers is
strongly encouraged to conduct basic research that will attract significant grant support
with little or no imperative for serving commodity groups through applied research.
This void could be addressed by reorienting the priorities of extension specialists.

Florida Entomologist 72(3)

September, 1989


An exciting prospect for improving resistance of trees to abiotic stress and biotic
agents may come through biotechnology. A portion of the gene responsible for toxin
production within Bacillus thuringiensis has been experimentally incorporated into the
DNA of tobacco plants that then produce the Bt toxin at insecticidal levels (Vaech et
al. 1987). This technology has exciting possibilities for both genetically adapting woody
plants to stressful urban conditions and providing plants with the genetic capacity for
auto-immunity against pests that threaten vitality or survival.


Anonymous. 1982. Genetic improvement and urban trees: A problem analysis for en-
vironmental forestry research pp. 510-519, in Anonymous [ed.], The Genetics
Working Group of the Consortium for Environmental Forestry Studies. U.S.
Govt. Printing Office: 65 pp.
BRIDGEMAN, P. 1979. Trees for town and country. David and Charles. London.
CARROW, J. R., AND K. GRAHAM. 1968. Nitrogen fertilization of the host tree and
population growth of the balsam woolly aphid, Adelges piceae (Homoptera: adel-
gidae). Canadian Entomol. 100: 478-485.
KIELBASO, J. J., AND M. K. KENNEDY. 1983. Urban forestry and entomology: A
current appraisal, pp. 423-440, in G. W. Frankie and C. S. Koehler [eds.] Urban
Entomology: Interdisciplinary perspective. Praeger Press.
LI, H. L. 1963. The origin and cultivation of shade and ornamental trees. Univ. of
Pennsylvania Press, Philadelphia.
PHILLIPS, L. F. 1979. Implementing a street tree inventory and planning system.
Weeds, Trees and Turf. 18(12): 18-23.
RAFFA, K. F. 1986. Devising pest management tactics based on plant defense
mechanisms, theoretical and practical considerations, pp. 301-327, in L. R.
Brattsten and S. Ahmad [eds]., Molecular aspects of insect-plant associations.
Plenum Press, New York.
SCHOENEWEISS, D. F. 1986. Water stress predisposition to disease. An overview,
pp. 157-174, in P. G. Ayres, [ed.] Water, Fungi, and Plants. Cambridge Univer-
sity, New York. 157-174 pp.
plants protected from insect attack. Nature. 328: 33-37.
WARING, R. H., AND G. B. PITMAN. 1983. Physiological stress in Lodgepole pine as
a precursor for mountain pine beetle attack. Z. Ang. Ent. 96: 265-270.
WRIGHT, J. W. 1976. Introduction to forest genetics. Academic Press, New York.

Cook: New Ecuadorian Damselfly


469 Crailhope Road
Center, Kentucky 42214
Research Associate, Florida State Collection of Arthropods,
Division of Plant Industry,
Florida Department of Agriculture and Consumer Services,
Gainesville, Florida 32602, USA.


Philogenia redunca sp. n., a new Ecuadoran megapodagrionid, is described and
illustrated from a series of 33 males and 25 females holotypee d: Yanamanaca, Napo
Province; allotype ?: Abitagua, Pastaza Province; both deposited in the Museum of
Zoology. Ann Arbor, USA). The male is distinguished by the shape of its anal appen-
dages. The female differs from all presently known species of Philogenia by possessing
a pair of lateral horns arising on the proepimeron and directed posteriorly. P. redunca
belongs in the cassandra group of species because the male of all six species possess a
peculiar meso-ventral flange-like process on the distal halves of their cerci. Within this
group redunca is most closely related to buenavista Bick & Bick and schmidti Ris.


Se describe e ilustra a Philogenia redunca sp. n., que es un megapodagrionido
Ecuatoriano, de una series de 33 machos y 25 hembras (holotipos 6: Yanamanaca, provin-
cia de Napo; alotipo 9: Abitagua, provincia de Pastaza; ambos depositados en el Museo
de Zoologia de Ann Arbor, E.U.A.). El macho se distingue por la forma de su ap6ndice
anal. La hembra difiere de todas las species conocidas de Philogenia porque posee un
par de cuernos laterales saliendo directamente posterior al proepimer6n. P. redunca
pertenece al grupo de species de cassandra porque los machos de las seis species
poseen un process peculiar meso-ventral parecido a un reborde en la mitad distal de su
cerci. Dentro de este grupo, redunca se relaciona mas a buenavista Bick & Bick y a
schmidti Ris.

Philogenia Selys is an exclusively Neotropical genus distributed from Costa Rica to
Bolivia. They are megapodagrionids of somber colors and secretive habits, at home in
deep forests where they may be encountered along rivulets and trails hanging on low
vegetation, and likely to be overlooked by all but the most keen-eyed observers.
Philogenia has received considerable attention in the literature. The genus was first
monographed more than half a century ago by Calvert (1924) who treated 17.species.
Westfall & Cumming (1956), Racenis (1959), and Dunkle (1986) together added a total
of 7 more. Bick & Bick (1988) have furnished the latest revision of the genus, adding
an additional 5 new species, and "species A" is discussed but not described. The purpose
of this paper is to described the Bick & Bick (1988) "species A".
This new species is unique by possessing structural specializations not present in
any other known species of the genus. For several years specimens of this form had
been in my collection, but they were badly broken and some important morphological
features were obliterated. It was thought best to try to obtain better examples before
describing them. When examining material for their revision of Philogenia, Bick & Bick
(1988) discovered a large series-of this species among the Ecuadoran material amassed


Florida Entomologist 72(3)

by the late C. H. Kennedy, and now held by the Museum of Zoology, University of
Michigan. Being aware of my prior recognition of this form as an undescribed species,
Dr. and Mrs. Bick very generously deferred describing the species to me and forwarded
the Kennedy material to me with their permission to include it in the type series.
ETYMOLOGY.-The specific name redunca (re-dun' ca) is from the L. "reduncus"=
"curved back, or hook-like" referring to the hook-shaped superior cerci of males in
dorsal view.

(Figs. 1-6)

Material examined.-All from ECUADOR.-HOLOTYPE male: Napo Province,
Yanamanaca, 6 October 1935, William Clarke-Macintyre (WCM) leg.-ALLOTYPE
female: Pastaza Province, Abitagua, 1,200 m, April 1937, WCM leg.-PARATYPES: 1
female, same data as holotype.-7 males, 6 females, same data as allotype.-2 males,
1 female, Morona-Santiago Province, Salado, Rio Upano, 1,550 m, (without date),
Lepoldo Gomez Alonzo (LGA) leg.-1 female, Napo Province, Rio Cotopino, February
1950, WCM leg.1 male, Napo Province, Rio Cotopino, March 1950, WCM leg.- 1 male,
Pastaza Province, Abitagua, 1,200 m, 4 June 1937, WCM leg.-1 male, 2 females,
Pastaza Province, Abitagua, 1,200 m, 26 May 1939, WCM leg.-1 female, Pastaza Pro-
vince, Abitagua, 1,000 m, 5 October 1939, WCM leg.- 1 female, Pastsza Province,
Abitagua, 1,100 m, 21 November 1939, WCM leg.-1 male, 4 females, Pastaza Prao-
vince, Abitagua, 1,100 m, January 1940, WCM leg.-1 male, Pastaza Province,
Abitigua, 1,100 m, 4 April 1940, WCM leg.-2 males, Pastaza Province, Abitagua, 1,100
m, 9 April 1941, WCM leg.-1 female, Pastaza Province, Abitagu, 1,100 m, 27 June
1940, WCM leg.-3 males, Pastaza Province, Abitigua, 1,000 m, 25 Febuary 1941,
WCM leg.-1 male Pastaza Provinc, Abitagua, 1,00 m, 1 March 1941, WCM leg.-1
female, Pastaza Province, Abitagua, "at mouth of the Guillermina Playa of Pastaza",
1,050 m, 16 May 1941, WCM leg.-1 female, Pastaza Praovince, Abitagua, "mouth of
Guillermina Creek", 1,050 m, 2 June 1941, WCM leg.-1 male, Pastaza Province,
Abitagua, 1,100 m, 4 June 1941, WCM leg.-1 female, Pastaza Province, Abitagua, "Rio
Pastaza Watershed", 24 October 1936, WCM leg.-1 male, Pastaza Province, "Pastaza
Watershed", December 1935, WCM leg.-1 female, Pastaza Province, "Rio Pastaza
Watershed near Mera", 1,200 m, 20 September 1937, WCM leg.-1 female, Tungurahua
Province, Banos, 17 August 1936, WCM leg.-2 males, Tungurahua Province, Rio Mar-
gajitas, 20 September 1937, WCM leg.-2 males, Morona-Santiago Province, Man-
gosisa, 850 m, (without date), LGA leg.-1 male, Tungurahua Province, San Francisco,
August 1938, (coll. unknown).-1 male, Tungurahua Praovince, San Francisco, 16 Oc-
tober 1937, (coll. unknown).-3 males, Tungurahua Province, Rio Topo, 1,400 m, June
1977, (coll. unknown).
Holotype male, allotype female, and paratypes collected by WCM and LGA (Ken-
nedy Collection) deposited in Museum of Zoology, University of Michigan, Ann Arbor,
USA. Remaining paratypes are in the collection of the author, to be deposited ultimately
in the Florida State Collection of Arthropods, Gainesville, USA.


Dimensions.-Total length 55.0 mm. Hindwing 37.5 mm. Abdomen (including ter-
minalia) 44.0 mm. Circi 1.7 mm.
Head.-Labium tan except darker on ligula and distal palps. Labrum yellow, nar-
rowly brown along anterior margin, a small brown spot in median depression. Genae
and bases of mandibles yellow, incisors brown. Anteclypeus obscure brownish, median
protuberance yellow. Postclypeus dull black. Frons and occiput dull black, with some


September, 1989

Cook: New Ecuadorian Damselfly

1 2


5 6
Figs. 1-6. Philogenia redunca spec. nov.: 1, dorsal view of male anal appendages;
2, lateral view of male anal appendages; 3, dorsal view of male prothorax; 4, lateral
view of female ovipositor; 5, dorsal view of female prothorax; 6, lateral view of female

mottled brown spots across vertex and occiput. Antennae with segment 1 and basal 2/3
of 2 pale, otherwise brown. Rear of head tan.
Prothorax.-Pronotum mostly tan, with pronounced longitudinal groove on meson;
anterior lobe mottled brown, its conspicuously upturned rim pale; posterior lobe brown

422 Florida Entomologist 72(3) September, 1989

with lateral tan spots, without a raised rim. Proepisternum and proepimeron dark
brown; near posterolateral angle is an obtuse pyramidal protuberance homologous to
the more pronounced horn-like process in the female.
Pterotrhorax.-Mesepisternum tan, with mid-dorsal carina, carinae along antealar
sinuses, and spot at upper end of humeral suture black. Mesepimeron, metepisternum,
and metepimeron tan, each with a central wide dark brown stripe. Undersurface of
metepimeron pale. Legs pale with brown carinae, setae, and tarsi. Prothoracic under-
surface, thoracic lateral dark stripes, coxae, and undersurface of metepimeron lightly
white pruinose.
Wings.-Veins dark brown. Pterostigma tan, 2.1 mm long, covering 4-5 cells in all
wings. Membrane slightly smoky, tips not opaque banded. Forewing postnodal cross-
veins 29-29, hindwing postnodal crossveins 25-26. M2 arises near 8th postnodal in fore-
wing, 7th in hindwing.
Abdomen.-Segment 1 widely pale on dorsum, dark brown ventrolaterally; segment
2 dark brown except for full length dorsolateral pale stripes, the ventrolateral dark area
lightly pruinose; segments 3-7 dark brown with pale basal rings, on 3 the pale extends
laterally as a 3/4 length narrow stripe; segments 8-10 dark brown to black, without pale
rings, densely encrusted with white pruinosity on entire dorsum of 9-10 and posterior
half of 8.
Anal appendages.-Cerci and paraprocts black. In dorsal view (Fig. 1) cerci forci-
pate, evenly curved inward, and dorsal and lateral surfaces thickly beset with acuminate
spines; distal halves produced into flange-like meso-ventral processes, these blunt, sub-
quadrate, and deflexed ventrad. Process furnished with a median transverse ridge, and
its proximal edge concave; in oblique view the flange-like process is auriculate ventrome-
dially, and entire cercus is concave internally. Length of cerci subequal to segment 9.
Paraproct (Figs. 1-2) wide at base, abruptly becoming narrow in distal half, the narrow
extremity curved dorsolaterally. Paraprocts equal ca. 0.80 the length of cerci.


Dimensions.-Total length 52 mm. Hindwing 37 mm. Abdomen (including ter-
minalia) 41 mm. Ovipositor 3 mm.
Head.-Colors and markings similar to male except labrum dark brown.
Prothorax and pterothorax.-Colors and markings as for male. Protuberance on
proepimeron (Fig. 6) more prominent than in male, developed as a robust posteriorly
directed horn.
Wings.-As for male except nodal index 29-30 / 27-27 for forewings and hindwings
Abdomen.-Colors and markings similar to male except lateral pale stripes more
extensive: 4/5 length on segments 3-4, 1/3 length on 5, 1/4 length on 6-7. Pruniosity on
dorsobasal 2/3 of 9 only.
Anal appendages.-Ovipositor (Fig. 4) extends 0.20 mm beyond tip of cerci, subequal
in length to segments 8 + 9. Cerci acute, pyramidal, subequal in length to segment 10.
Styli slightly longer than cerci (by a ratio of 5 to 4). Epiproct in lateral view 1/4 length
of cerci.


Bick & Bick (1988) in their revision recognized six infrageneric categories within
Philogenia which they have referred to as "species groups". These groups were estab-
lished on the basis of structural similarities, particularly the shape of male anal appen-
dages. The male of redunca, by reason of the similarity of its anal appendages, seems

Cook: New Ecuadorian Damselfly

most closely related to buenavista Bick & Bick and schmidti Ris. These, along with the
slightly modified cassandra Hagen, polyxena Calvert, and umbrosa Ris, form the "Cas-
sandra Group", of the above authors. The female of redunca because of its specialized
prothoracic horns stands apart from all other presently known females in the genus.
Species identification in Philogenia has been based chiefly on male morphological
characteristics in most previous papers. The male anal appendages show a wide range
of diversity among the different species and are highly diagnostic for species recogni-
tion. Wing coloration (dark bands present or absent) and color patterns of the body are
also useful characters. To date, no one seems to have attempted utilizing penis morphol-
ogy for taxonmic classification of the species in this genus, and examples of different
species in my possession are too meager to indicate if useful penis characters do exist.
Prothoracic morphological features are traditionally the most useful characters to
separate females of megapodagrionids, but, especially in Philogenia, difficulty is fre-
quently experienced in correctly associating females with their male counterparts. All
of the approximately 200 species examined in this study, were collected by local insect
sellers who provided only minimum collecting data, and no data at all to indicate if any
were taken in copula, or otherwise to suggest the possible association of sexes. Several
previous studies have not treated Philogenia females because of the supposed paucity
of annectant characters between sexes. This has resulted in the male sex only being
described for several species. The importance of correctly associating the sexes of re-
dunca was realized in the beginning of this study because the "feature story would
be the aberrant prothoracic horns of the female. Very fortunately this specialized
character was found to be homologous in both sexes, and though less pronounced in the
male, it has provided the positive annectant character needed for associating the sexes
of this species.
My studies have also revealed the existence of heretofore unrecognized prothoracic
morphological structures in both sexes. I believe these provide useful diagnostic charac-
ters for determining females and associating them with their correct male counterparts
in the genus. These characters were found to be consistently distinctive in the males of
10 species and females of 7 species of this genus available for study.
The characters referred to are distinctive arrangements of ridges and depressions
on laterotergal surfaces of prothorax and shape of its anterior, lateroventral, and pos-
terior margins (which may be entire, or scalloped in various patterns, in different
species). These newly, recognized characters have been utilized to associate several
species of previously unrecognized females with their described male counterparts.
Preparation of a paper describing them is planned.
Nothing is known concerning the biology or behavior of P. redunca, but prothoracic
accessory appendages in other Odonata have been shown to have various functional
roles in the tandem mating process. They may function as tactile sensory organs to aid
species recognition, as well as acting as part of the actual mechanics of the coupling
process. The absence of similar specialized prothoracic accessory appendages in other
species of Philogenia is a peculiar anomaly not well understood. The trait of strong
similarities in male anal appendages among species of Bick & Bick's "Cassandra Group"
probably was developed in a very early ancestral form. If specialized prothoracic horns
had developed in parallel with the male's flange configured anal appendages as a compat-
ible prothoracic configuration, some degree of evolutionary development of horns would
be expected to occur in other females belonging to the "species group" in question. That
redunca developed its own unique prothoracic configuration suggests its comparatively
recent appearance, and after isolation from other congeners in the "Cassangra Group".
A possible hypothesis for evolutionary development of continental species groups post-
ulate the continuous distribution of single ancestral forms from Columbia to Bolivia.
Subsequent uplift of the Andes Mountain chain would form barriers across the continu-


Florida Entomologist 72(3)

September, 1989

ous distribution of these ancestral forms causing their division into disjunct populations
from which the present species groups have evolved.
P. redunca is the most specialized species which has yet evolved in the "Cassandra
Group" and has developed individual characteristics suitable for prosperity in its par-
ticular niche.


I am very grateful to Dr. George Bick and Mrs. Juanda Bick, who with characteris-
taic generosity permitted me to include material in this study which they had discov-
ered, I also thank both of them for suggestions and guidance during the study. I thank
Mr. Mark F. O'Brien and the Insect Division, Museum of Zoology, University of Michi-
gan, for the opportunity to examine the Kennedy material of the new species, and also
for the opportunity to study an additional large lot of Philogenia females. I thank Dr.
S.. W. Dunkle, Dr. H. V. Weems, Jr., and Dr. M. J. Westfall, Jr., for reviewing the
Contribution No. 705, Bureau of Entomology, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville, FL 32602, USA.


BICK, G. H. AND J. C. BICK, 1988. A review of males of the genus Philogenia, with
descriptions of five new species from South America (Zygoptera: Megapod-
rionidae). Odonatologica 17: 9-32.
CALVERT, P. P. 1924. The generic characters and the species of Philogenia Selys
(Odonata: Agrionidae). Trans. Am. ent. Soc. 50: 1-56.
DAVIES, D. A. L. AND P. TOBIN. 1984. The dragonflies of the world. Vol. 1 Soc. int.
odonatol. rapid Comm. (Suppl.) 3: 1-127.
DUNKLE, S. W., 1986. Four new species of Philogenia damselflies from Ecuador and
Colombia (Zygoptera: Megapodagrionidae). Odonatologica 15: 43-50.
HAGEN, H. A., 1869. Zur Odonaten-Fauna von Neu-Granada nach Lindigs
Sammlungen. Stett. ent. Zeitg. 30: 256-263.
MAY, M. L., 1898. Status of Philogenia leonora Westfall & Cumming (Zygoptera:
Megapodagrionidae). Odonatologica 18: 95-97.
RACENIS, J., 1959. Notas taxonomicas sobre la familiar Megapodagrionidae (Odonata:
Zygoptera) con la sinopsis de la species Venezolanas. Acta Biol. Venezuelica 2:
Ris, F., 1918. Libellen (Odonata) aus der Region der amerkanischen Kordilleren von
Costarica bis Catamarca. Arch. Naturgesch. (A) 82(9): 1-197.
SELYS LONGCHAMPS, E. de, 1862. Synopsis des Agrionines, Troisieme legion;
Podagrion. Bull. Acad. R. Belge (2) 14: 3-42.
SELYS LONGCHAMPS, E. de, 1886. Revision du Synopsis des Agrionines, Premiere
Parte; comprenant les legions Pseudostigma, Podagrion, Platycnemis et Pro-
tonevra. Mem. Cour. Acad. R. Sci. Belgique 38: 1-223.
WESTFALL, M. J. AND R. B. CUMMING 1956. Two new species of Philogenia from
the Panama Canal Zone (Odonata: Coenagriidae). Bull. Florida State Mus. 1:

Donnelly: New Honduran Damselfly


2091 Partridge Lane, Binghamton, NY 13903
Research Associate, Florida State Collection of Arthropods,
Division of Plant Industry,
Florida Department of Agriculture and Consumer Services,
Gainesville, Florida 32602


Philogenia strigilis n. sp. from northwestern Honduras extends the range of the
genus more than 600 km to the north, the male has a remarkably simple superior
appendage unlike any other species in Central America, but reminiscent of P. boliviana
from Bolivia. The male inferior appendage and the female hind lobe of the prothorax
both resemble these structures on P. augusti from Panama.


Philogenia strigilis n. sp. de la parte noraeste de Honduras alarga la distribuci6n
del g6nero mas que 600 kil6metros al norte. El macho tiene un ap6ndice superior muy
sencillo que no se semeja a ap6ndices de otras species de Centro AmBrica, pero es
semejante a P. boliviana de Bolivia. El ap6ndice inferior del macho y el 16bulo posterior
del prot6rax de la hembra son semejantes a ellos de P. augusti de Panama.

The discovery of a new species of Philogenia from northwestern Honduras extends
the range of the genus northward from northern Costa Rica a distance of at least 600
km. The new species is possibly an aberrant member of the carrillica group most closely
related to the Panamanian species augusti.

Philogenia strigilis Donnelly, n. sp. (Figs. 1-4)

HOLOTYPE MALE. Head: dorsally dark brown; labrum, genae, and sides of mandibles
bright creamy blue-green, grading to creamy yellow; thin dark line along distal margin
of labrum. Dorsum of head dark with obscure lateral pale marks, centrally constricted,
between eyes and lateral ocelli; occiput with very obscure pale markings.
Thorax: dark, pruinescent on sides; prothorax obscure pale brown on posterior part
of mid lobes and lateral part of hind lobe; hind lobe erect, with obtuse angulation on
lateral corner. Pterothorax: mesepisternum dark with obscure thin pale anterior border
extending posteriorly along antehumeral suture 4/5 length of sclerite; sides of thorax
dark with broad, obscure pale bands on 1st and 2nd lateral sutures extending anteriorly
to mesinfraepisternum and metinfraepisternum; metepimeron centrally dark with broad
pale border; metasternum pale. Wings: venation dark red brown; pterostigma yellow
brown, 3.2 mm long; wing tips very slightly infumated. Legs: fore legs obscurely dark,
mid and hind legs pale, with dorsal surface of femora and tibiae obscurely dark.
Abdomen: dark, pruinescence on sides of 1 and 2; obscure yellow lateral dash on 2;
basal yellow ring obscurely connected dorsally on 3-7 and short basal-lateral thin yellow
lines on 3-5, decreasing in size posteriorly. Superior appendages: simple, decurved,
twisted, with concave ventral surface in apical 1/3 visible in lateral view; no distinct

Florida Entomologist 72(3)

1 2

Figs. 1-4 Philogenia strigilis. Figs. 1-3, lateral, apical, and ventral views of the male
terminal appendages. Fig. 4, antero-dorsal view of the hind lobe of the prothorax and
the mesostigmal laminae of the female.

meso-ventral process sensu Bick and Bick; in dorsal view with small basal-mesal exten-
sion terminating in a low mesal rounded tubercle; sinuous dorsal-mesal edge thickened
and denticulate; dorsal surface covered with short spines. Inferior appendages long,
divaricate in ventral view, with apices straight, rounded at tip, and slightly curved
dorso-mesally; with dorso-mesal margin continuing mesally to a prominent dorsally
projecting spine; basal-lateral part with a low apically pointed angulated projection.
ALLOTYPE FEMALE. Colored as in male. Head: obscure pale marks on dorsum of head
more extensive, coalesced to form a transverse band through ocelli; obscure marks on
occiput more extensive, including obscure round spots behind eyes.
Thorax: hind lobe of pronotum (Fig. 4) terminates laterally in rounded, protuberant,
slightly inflated lobes; thorax mainly pale brown (above) grading to obscure yellow on
sides; mesepimeron and mesinfraepisternum dark, pale band on 1st lateral suture ex-
tending to dorsal 2/3 of metepisternum, whose ventral 1/3 is dark; metepimeron with
dark center broadly bordered by pale; metasternum pale. Wings: as in male. Legs: as
in male but dorsal surfaces of mid and hind femora paler.
Abdomen: dark brown, with thin yellow lateral line on 2 to basal 4/5 of 3 and 4, 3/4
of 5 and 6, and 1/2 of 7, becoming more obscure posteriorly; obscure basal yellow rings,

September, 1989

Donnelly: New Honduran Damselfly

interrupted dorsally, on 3 7; 9 with paired subbasal dorsal round pale spots; ventral
margins of tergites 3 7 pale, with color expanding posteriorly; ovipositor extending
slightly beyond tip of 10, valves with finely rugose surface.
Dimensions: Holotype male: abdomen 44 mm; hind wing 38 mm. Allotype female:
abdomen 42 mm; hind wing 37.5 mm.
VARIATIONS WITHIN TYPE SERIES. The holotype and 9 paratype males range in size
as follows: abdomen 43 51 mm; hind wing 36.5 42 mm. There is a slight range in the
extent of pale color on the top of the head. Darker specimens have the dorso-lateral
spots divided centrally into two spots on each side; paler specimens have these spots
virtually unconstricted and have, in addition, more prominent obscure pale spots on the
MATERIAL STUDIED. All specimens: HONDURAS: Cortes; Rio Piedras at San Pedro
Sula, 16 July 1971. This stream is in a protected, forested watershed for the city and
yielded in 1971 a rich selection of Odonata, including Heteragrion eboratum Donnelly,
Archilestes latialatus Donnelly, an undescribed Palaemnema, and numerous species of
Argia and Hetaerina. I visited the same locality in 1976 following a major tropical
storm. At that visit, the vegetation bordering the stream had been thoroughly de-
stroyed by flooding, leaving a small, clean stream flowing in the center of a sun-baked
broad avenue of cobbles and boulders. As I expected, the formerly rich zygoptera fauna
was essentially absent, although numerous pseudostigmatids were seen in the surround-
ing more elevated forest.
The holotype and allotype are deposited in the Florida State Collection of Ar-
thropods, Gainesville.
ETYMOLOGY. The name is that of a Roman flesh-scraping implement, which bears a
strong resemblance to the distinctive male superior appendage.


Bick and Bick (1988) published the first attempt to group the species of this complex
genus, basing their categories primarily on the male superior appendage and secondarily
on the inferior. P. strigilis has a remarkable simple decurved and twisted superior
appendage which lacks a distinct meso-ventral process; except for its smaller size, it
bears a strong resemblance to that of P. boliviana. The Bicks concluded that this simple
superior appendage implied a primitive position within the genus. If the resemblance
between the superior appendages of these two species is meaningful, and if the Bick's
conclusion that this form is primitive is accepted, then we might conclude that the two
most primitive species lie at the opposite extremes of geographic range.
The inferior appendage of P. strigilis, however, is very much unlike that of
boliviana. Its prominent subbasal, dorsally directed spine is unlike that of any other
species of the genus, except perhaps for augusti, which it resembles (Calvert's descrip-
tion of augusti omits mention of this spine, and it is barely visible in his figures). The
inferior appendage also bears a resemblance to three other Central American species
with widely differing superior appendages: carrillica, leonora, and champion. How-
ever, each of these lacks the dorsal-directed spine terminating the dorso-mesal margin
of the inferior appendage. Bick and Bick placed each of these four species in his champ-
ioni group (not separating leonora and champion, May, 1989). I believe the broad
resemblance of the inferior appendages places all of these species in the same group,
but the very simplified superior appendage of strigilis suggests that the character of
this appendage should not be the primary character for grouping species within this
The female prothorax of strigilis, with its prominent lateral lobes, is most like that
of augusti among the relatively few species whose females are known.


428 Florida Entomologist 72(3) September, 1989

I conclude that, at least for the Central American species of the genus with which
I am familiar, P. strigilis is an aberrant member of the carrillica group, and I would
caution that the superior appendage may be a deceptively plastic appendage which
should be used for phylogenetic groupings only with great care.
I have taken a single female of Philogenia near Puerto Barrios, Guatemala, which
is about 75 km WNW of San Pedro Sula. This female resembles strigilis but has a
darker face and a simply rounded lateral margin of the hind lobe. Unless strigilis is
strongly polymorphic (Calvert, 1924, was puzzled by apparent polymorphism in females
of carrillica Calvert.), then this female belongs to yet another species.


I am most grateful to George Bick for his helpful and substantive comments on the
first draft of this paper.
Contribution No. 711, Bureau of Entomology, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville, Florida 32602.


BICK, G. H. AND BICK, J. C. 1988. A review of the males of Philogenia, with descrip-
tions of five new species from South America (Zygoptera: Megapodagrionidae).
Odonatologica 17(1): 9-32.
CALVERT, P. P. 1924. The generic characters and species of Philogenia Selys
(Odonata: Agrionidae). Trans. Am. Ent. Soc. 50: 1-56.
MAY, M. L. 1989. Status of Philogenia leonora Westfall & Cumming (Zygoptera:
Megapodagrionidae). Odonatologica 17: 95-97.


2091 Partridge Lane, Binghamton, NY 13903
Research Associate, Florida State Collection of Arthropods,
Division of Plant Industry,
Florida Department of Agriculture and Consumer Services,
Gainesville, Florida 32602


The genus Epigomphus Selys 1854 in Mexico and northern Central America contains
six described and three new species. E. maya n. sp. from the Maya Mountains of Belize
is closest to E. quadracies from western Guatemala, Costa Rica, and Panama. E. flinti
n. sp. from Oaxaca and E. sulcatistyla n. sp. (originally placed with E. paulsoni from
Veracruz) are closest to E. paulsoni, which is limited to Chiapas. Males of Epigomphus
are separated most reliability by the form of their abdominal appendages.


El genero Epigomphus Selys 1854 en Mexico y la parte norte de Centro America

Donnelly: Odonata from Belize & Mexico

contiene seis esp6cies descritas y tres nuevas. E. maya n. sp. de las "Maya Mountains"
de Belize esta relacionada con E. quadracies de la parte occidental de Guatemala, Costa
Rica, y PanamA. E. flinti n. sp. de Oaxaca y E. sulcatistyla n. sp. (originalmente puesta
con E. paulsoni) de Veracruz estan relacionadas con E. paulsoni, que esta limitada a
Chiapas. Machos de Epigomphus se diferencian con mAs seguridad por la forma de los
apendices abdominales.

Although there are 9 species of Epigomphus in northern Central America and
M6xico, only one of these (subobtusus Selys, 1878) is widespread (Costa Rica to Mexico)
and represented in collections by numerous specimens. The remainder are rarely taken
and apparently very restricted in their occurrences. The holotype of E. quadracies
(Calvert, 1903) was taken in western Guatemala, but no subsequent specimens have
been taken north of Costa Rica. E. paulsoni (Belle, 1981) was described from 4 males,
2 from Veracruz and 2 from Chiapas. The Veracruz specimens are here placed in a
separate species: E. sulcatistyla. E. crepidus (Kennedy, 1936) is known from a few
males from Nayarit and Chiapas. E. clavatus (Belle, 1980) is known from a single male
from Alta Verapaz, Guatemala. E. donnellyi (Gonzalez-Soriano & Cook, 1988) is known
from about a dozen specimens from southeastern Veracruz. Three new species are
described herein: E. flinti from a single locality in Oaxaca, E. sulcatistyla from a single
locality in Veracruz, and E. maya from the Maya Mountains of Belize.
The present study shows that certain characters used in the gomphidae are of ques-
tionable value in separating these species of Epigomphus. The size, color pattern, wing
venation, and hamules of the 2nd segment are all very similar. All of these species have
tiny spines on the dorsum of the 10th abdominal segment denticless of Belle, 1980, 1981);
however, these seem to have limited diagnostic significance.
For the males the terminal abdominal appendages are highly differentiated and
appear to allow groupings of species. I introduce here a brief terminology to label the
protuberances of the dorsal surface of the inferior appendage. The apex (A) of each
ramus is generally a distinct point; in some species it is either indistinctly separated
from the 1st spine (Sl) (sulcatistyla; maya) or fused with this spine (paulsoni). The
1st, or sublateral, spine (Sl) is generally subapical and is especially distinct in lateral
view. The 2nd, or submesal, spine (S2) is less visible than Sl in lateral view in four
species (clavatus, sulcatistyla, paulsoni, flinti) and absent in the remaining 3. The
tubercle (T) is the apical end of a rounded submesal ridge on the ramus. In four species
it protrudes beyond the margin of the ramus (maya, quadracies, paulsoni, flinti). In
sulcatistyla it has a prominent subapical swelling.
The females of only 3 species are known, not including any of the new species, and
I am unable to provide any means for the potential differentiation of the unknown

Epigomphus maya Donnelly, new species (Figs. 1, 5, 9)

HOLOTYPE MALE. Head: red brown, pale olive green as follows: rounded lateral
spots on labrum, ventro-lateral dashes on postclypeus, transverse band on frons, sides
of mandibles, and genae; yellow half circles around outer border of lateral ocelli. Crest
of occiput with a very shallow concave angulation; shallow indentations at sides of
Thorax: prothorax pale brown, obscurely mottled, with paired yellow spots on hind
margin of middle lobe. Pterothorax red brown above, obscurely pale yellow below,
yellow stripes as follows: 1st and 2nd on mesepisternum, the 2nd (antehumeral) narrow
and expanded posteriorly to a rounded spot; 3rd on mesepimeron and 1/4 its width, with


Florida Entomologist 72(3)


Figs. 1-4. Lateral view of male terminal appendages. Fig. 1, E. maya. Fig. 2, E.
quadracies. Fig. 3, E. clavatus. Fig. 4, E. crepidus.

expanded, quadrate posterior end; 4th on metepisternum and 1/2 its width, with round,
dorsal excavation on posterior 1/2; 5th on metepimeron and 1/2 its width, with expanded,
rounded posterior end; transverse isolated bars on anterior edge of mesepisternum;
metasternum pale. Wings: venation and pterostigma black; thickened antenodals 6/7 on
fore wings, 6/7 on hind wings; 3 cells between Al and A2, and between A2 and A3 and
the hind wing margin; 3 cells between base of wing and A3. Legs: femora pale with
darker exterior apical streaks on fore ad mid femora; tibiae and tarsi dark.
Abdomen: dark brown dorsally, sides greenish on 1 to 3, pale on anterior 1/4 (to
transverse carina) of 4 to 6; 7 and 8 largely yellow, obscurely brown on posterior 1/4 of
7 and posterior 1/2 of 8; 9 dark; 10 reddish brown; thin yellow dorsal line on 1 to 4; small
dorso-apical spot on 5 and 6. Terminal appendages: red brown, tips black; superior
thick, decurved (Fig. 1), with truncate tip and thickened apical edge with 7 inferior
denticles, angle of tip 350 from midline viewed ventrally (Fig. 9); inferior appendage
shorter than superior, shallowly forked with deep, rounded mesal sulcus and sharp
dorso-apical projection directed cephalad (Sl of Fig. 5), short apex (A), and submesal
dorsally directed tubercle (T) on inner edge of fork. Genitalia of 2nd segment: anterior

September, 1989

Donnelly: Odonata from Belize & Mexico 431

hamule thin, flat, with round, tapering tip; posterior subcylindrical, tapering apically
to rounded tip, with short black denticles on anterior surface.
Dimensions: abdomen 41 mm; hind wing 35 mm.
VARIATIONS AMONG PARATYPE SERIES. Two paratype males have the abdomen 39
and 41 mm, and the hind wing 34 mm. One paratype male has segment 8 totally dark
on the dorsum and the other dark in the apical 1/2 of the dorsum.
MATERIAL EXAMINED. Holotype, BELIZE: Cayo Dist.; Mountain Pine Ridge, Little
Vaqueros Cr., Chiquibul Rd. (17 03 N, 880 57' W; 500 m), 22-25 July 1983, Coll. T.
Donnelly. One paratype has the same data; the other s from a nearby stream on Wolfson
Drive (160 55' N, 880 53' W; 600 m), 23 July 1983, Coll. T. Donnelly. The holotype is
deposited in the Florida State Collection of Arthropods, Gainesville.
The streams in which these insects live are small, clean, probably permanent streams
in the dominantly pine forest of the Maya Mountains. The first flows on coarse-grained
granite and has a coarse sand to gravel bottom with minor mud; the second flows on
sedimentary rocks and has a mud to sand bottom.
ETYMOLOGY: The name is that of the isolated mountain range where the species occurs.
This species is the only Epigomphus that has been taken in Belize. The form of its
inferior appendage is similar to that of quadracies, especially in the appearance of the
median sulcus and ornamentation (A, Si, and T; in both species S2 is apparently mis-
sing). The superior appendage is of generalized form and resembles several diverse
species: crepidus (Fig. 4), clavatus (Fig. 3), paulsoni (Fig. 14), sulcatistyla (Fig. 13),
and even the very different subobtusus (not shown).

Epigomphus flinti Donnelly, new species (Figs. 15, 18, 21, 24)

HOLOTYPE MALE. Head: red brown, pale olive green as follows: large rounded lateral
spots on labrum (which also has a vividly black rim), ventro-lateral dashes on post-
clypeus, transvers centrally interrupted band on frons, sides of mandibles and genae;
vertex completely dark; crest of occiput with shallow lateral concave excavations.
Thorax: prothorax obscure red brown, with paired yellow lateral and paired mesal
spots on the hind margin of middle lobe; hind lobe dark brown. Pterothorax: red brown
above, obscurely pale below, yellow stripes as follows: pale anterior margin of mesepis-
ternum not connected with thin 1st stripe, the 2nd (antehumeral) reduced to a thread
and expanded posteriorly to a rounded spot; 3rd on mesepimeron and 1/6 its width, with
flared posterior end; 4th on metepisternum and 1/2 its width, with round dorsal excava-
tion on posterior 1/2; 5th on metepimeron and 2/5 its width, with expanded, rounded
posterior end; transverse isolated bars on anterior edge of mesepisternum; metasternum
pale. Wings: venation black, pterostigma red brown. Thickened antenodals in position
7/7 on fore wings, 6/7 on hind wings; 3 cells between anal vein and hind wing margin,
in space between Al and A2, and between A2 and A3; 3 cells between base of wing and
A3. Legs: femora pale, darkened apically; tibiae and tarsi dark.
Abdomen: dark brown dorsally, sides greenish on 1, 2, and anterior 5/6 of 3; pale on
anterior 1/4 (to transverse carina) of 4 to 6; basal half of 7 yellow, extended posteriorly
on sides; 8 and 9 dark brown; 10 red brown; thin yellow dorsal line on 1 to 4; small
dorso-apical spot on 5. Terminal appendages: dark brown; superior thick, decurved,
very slightly apically excavated in lateral view (Fig. 15), with truncate tip and thickened
apical edge bearing 2 large teeth and a few denticles; angle of tip 300 from mid line
viewed ventrally (Fig. 24); inferior appendage shorter than superior, shallowly forked,
with small apical point (A of Figs. 18, 21), largest dorsal spine (S1) flattened, subapical
on outer edge, small submesal spine (S2) and low mesal paired tubercles (T), with T,
S2, and S1 lying on straight line on each ramus. Genitalia of 2nd segment: anterior
hamule thin, flat, with rounded, tapering tip; posterior hamule flat, tapering apically to
rounded tip, with short, black denticles on anterior surface.

Florida Entomologist 72(3)





i Q








Figs. 5-12. Dorsal view (Figs. 5-8) and ventral view (Figs. 9-12) of male inferior
appendages. A = apex; S1 = subapical spine; S2 = submesal spine; T = mesal tubercle.
Fig. 5, E. maya (S2 not present). Fig. 6, E. quadracies (S2 not present). Fig. 7, E.
clavatus. Fig. 8, E. crepidus (S2 not present). Fig. 9, E. maya. Fig. 10, E. quadracies.
Fig. 11, E. clavatus. Fig. 12, E. crepidus.

Dimensions: abdomen 42.5 mm; hind wing 35 mm.
VARIATIONS AMONG PARATYPE SERIES. Of the 6 paratype males, 2 have the 2nd


September, 1989

Donnelly: Odonata from Belize & Mexico 433

thoracic stripe discontinuous. The abdomens of the series range from 39 to 42.5 mm and
the hind wings from 32 to 35 mm.
MATERIAL EXAMINED. Holotype and 6 paratype males, MEXICO: Oaxaca, 8 km S.
of Valle Nacional, 25 May, 1981, Coll. C. M. and O. S. Flint, Jr. The holotype and most
of the paratypes are deposited in the U.S. National Museum, Washington. One paratype
is deposited in the Florida State Collection of Arthropods, Gainesville.
ETYMOLOGY: The species is named for its collector, Ollie Flint. The dedication to him
also reflects the many extensive and valuable Odonata collections he has made while
pursuing trichoptera throughout the New World, as well as his generous and unfailing
assistance to odonatologists.
This species is close to E. paulsoni and E. sulcatistyla, whose ranges straddle the
occurrence of flinti. The 3 species are distinguished by (1) the tips of the superior
appendage in dorsal or ventral view (Figs. 13-18), which are truncated at a far more
acute angle in flinti than in paulsoni or sulcatistyla; and (2) the form of the inferior
appendage. In flinti the subapical spines S1 and S2 of each ramus are rounded in
section, pointed, and subequal in length (Fig. 18). They give the appearance of a pair
of horns on each ramus, in contrast to the flattened S1 and blunt S2 of the other 2
species (Figs. 16, 17).

Epigomphus sulcatistyla Donnelly, new species (Figs. 13, 16, 19, 22)

Epigomphus paulsoni: Belle 1981:61 (in part)

HOLOTYPE MALE. Head: red brown, pale olive green as follows: large rounded lateral
spots on labrum (which also has a narrowly black rim), ventro-lateral dashes on post-
clypeus, transverse band on frons, sides of mandibles and genae; vertex completely
dark; crest of occiput with shallow lateral concave excavations.
Thorax: prothorax obscure pale brown, with middle lobe pale laterally and with
paired yellow mesal spots on the hind margin; hind lobe dark brown. Pterothorax: red
brown above, obscurely pale below, yellow stripes as follows: pale anterior margin of
mesepisternum not connected with thin 1st stripe, the 2nd (antehumeral) thin and sub-
equal in width throughout; 3rd on mesepimeron and 1/5 its width, with expanded post-
erior end; 4th on metepisternum and 1/2 its width, narrowed and then expanded post-
eriorly into a circular spot; 5th on metepimeron and 1/2 its width; transverse isolated
bars on anterior edge of mesepisternum; metasternum pale. Wings: venation black,
pterostigma dark red brown. Thickened antenodals in position 6/7 on fore wings, 5/6 on
hind wings; 3 cells between anal vein and hind wing margin, in space between Al and
A2, and between A2 and A3; 3 cells between base of wing and A3. Legs: femora pale,
darkened apically; tibiae and tarsi dark.
Abdomen: dark brown dorsally, sides heavily marked with greenish on 1 and 2,
obscure lateral pale dashes on 3 (5/6 length) to 6 (1/2 length); small pale dorsolateral
basal lines on 3 to 6, shortened posteriorly to spots; 7 largely pale, obscurely darker
brown on dorsoposterior margin; 8 and 9 obscurely brown; 10 dark red brown. Terminal
appendages: dark red brown; superior thick, decurved, tips infolded inwards (Fig. 13),
apical-ventral margin bearing dark teeth; angle of tip 850 from mid line viewed ventrally
(Fig. 16); inferior appendage shorter than superior, forked, with deep mesal sulcus;
apex (A) fused with flattened sublateral spine (Sl); submesal spine (S2) midway between
S1 and T; tubercle (T) produced subapically into high and prominent submarginal swel-
ling (Figs. 16, 19). Genitalia of 2nd segment: anterior hamule thin, flat, with rounded,
tapering tip; posterior hamule subcylindrical-triangular, tapering apically to rounded
tip, with short, black denticles on anterior surface.
Dimensions: abdomen 41 mm; hind wing 34 mm.

Florida Entomologist 72(3)



Figs. 13-15. Lateral views of male terminal appendages. Figs. 13, E. sulcatistyla.
Figs. 14, E. paulsoni. Figs. 15, E. flinti.

VARIATIONS AMONG PARATYPE SERIES. The paratype male (figured in part as the
paratype of E. paulsoni Belle, 1981) has the abdomen broken and the hind wing 34.5
mm. There are no important differences between the two specimens.
MATERIAL EXAMINED. Holotype and paratype males, MEXICO: Veracruz; stream at
Coyame (1300'), 1 July 1965, coll. D. R. Paulson. The holotype is deposited in the
Florida State Collection of Arthropods, Gainesville, and the paratype in the Paulson

s< 17


18 A
T, S2 S



Figs. 19-24. Inclined dorsal-lateral (Figs. 16-18), dorsal (Figs. 19-21), and ventral
(Figs. 22-24) views of male terminal appendage. S1, S2, T, and A as for Figs. 5-8. Figs.
16, 19, 22, E. sulcatistyla (T not labeled). Figs. 17, 20, 23, E. paulsoni. Figs. 18, 21,
24 E. flinti.

September, 1989

Donnelly: Odonata from Belize & Mexico

Belle (1981) discussed the differences between the Veracruz and Chiapas specimens
at some length in the original description of E. paulsoni. However, he did not dicsuss
the very different inferior appendages, which are the crux of the present description.
The discovery of E. flinti intermediate in range between the other 2 forms forces us to
focus on the morphological differences among the members of this group of sibling


The 9 species of Epigomphus of Guatemala, Belize, and M6xico are the most north-
erly in the genus and form a relatively homogeneous group within a total of 22 species
extending south to Brazil and Argentina. These 9 species fall into 3 groups of 2 species
each and a group of 3 species. The first pair of species are subobtusus and donnellyi,
which are further closely related to E. westfalli (Donnelly, 1985) from Nicaragua. These
species are not figured here, but have been figured recently in Donnelly (1985) and
GonzAlez-Soriano and Cook (1988). They are easily distinguished from the remaining 7
by their long and deeply divaricate inferior appendage. Of the remaining species, maya
and quadracies have similar inferior appendages, both of which have diminished apices
(A of Figs. 5, 6) and a prominent median sulcus. The species crepidus and clavatus have
elongate apices (A of Figs. 7, 8) and diminished mesal tubercles (T), with different
arrangements of spines (S1 and S2, the latter absent in crepidus). The species flinti,
sulcatistyla, and paulsoni have short apices (A of Figs. 19-21) set very close to spines
S1, and spines S2 similarly developed. The differences among these 3 species have been
noted above.


I am very grateful to D. Paulson for the loan of a paratype of E. paulsoni and a
specimen of e. crepidus, and the provision of the holotype of E. sulcatistyla; to J. Belle
for the loan of a paratype of E. paulsoni which has become the paratype of E. sulcatis-
tyla; to G. Jurzitza for the loan of the holotype of E. clavatus; and to R. Garrison for
the loan of additional specimens. G. Bick, M. Westfall, and S. Dunkle made valuable
criticisms of an earlier draft of this manuscript.
Contribution No. 701, Bureau of Entomology, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville, Florida 32602.


BELLE, J. 1980. A new species of Epigomphus from Guatemala (Odonata: Gom-
phidae). Ent. Berichten 40: 136-138.
BELLE, J. 1981. A new species of Epigomphus from Mexico (Odonata: Gomphidae).
Ent. Berichten 41: 61-63.
CALVERT, P. E. 1903. On some American gomphinae (Odonata). Ent. News 14: 183-
DONNELLY, T. W. 1985. Epigomphus westfalli sp. nov., a new dragonfly from
Nicaragua (Odonata: Gomphidae). Odonatologica 15(1): 37-41.
GONZALEZ-SORIANO, E. AND COOK, C. 1988. Una nueva especie de Epigomphus
Selys 1854 (Odonata: Gomphidae) del estado de Veracruz, M6xico. Folia Ent.
Mexicana No. 74: 5-12.
KENNEDY, C. H. 1936. Epigomphus crepidus a new dragonfly (Odonata: Gomphidae)
from Nayarit, Mexico with notes on the genus. Ann. Ent. Soc. Amer. 29: 126-
SELYS LONGCHAMPS, E. de 1878. Quatri6mes additions au synopsis des gomphines.
Bull. Acad. Royale de Belgique (2) 46: 468-571 (original pagination).


436 Florida Entomologist 72(3) September, 1989


2091 Partridge Lane, Binghamton, NY 13901 USA
Research Associate, Florida State Collection of Arthropods,
Division of Plant Industry
Florida Department of Agriculture and Consumer Services
Gainesville, Florida 32602

Protoneura sulfurata n. sp., from the Caribbean coast of Costa Rica, differs from
P. aurantiaca in its bright sulfur-yellow color of the male and in several minor structural
details. The two species belong to a series (including P. amatoria and cupida) that
originated in South America and has reached M6xico. A second group of species (P.
tenuis, calverti, ailsa, viridis, capillaris, and probably cara) originated in South
America, occupied the Antilles, and has reached northern Central America, Mexico,
and the United States. A third group (which includes the three somewhat aberrant
species P. peramans, corculum, and sanguinipes) probably originated in South America
and has reached Central America and the Greater Antilles. The most derived species
are found at the extreme geographic limits of each group.

Protoneura sulfurata n. sp. de la costa del Mar Caribe de Costa Rica se diferencia
de P. aurantiaca por el color azufre-amarillo del macho y por diversos detalles estruc-
turales. Las dos species pertenecen a un series (incluyenda P. amatoria y cupida) que
origin en Sur America y ha alcanzado a M6xico. Un segundo grupo de species (P.
tenuris, calverti ailsa, viridis, capillaris, y probablemente cara) origin6 en Sur
America, ocup6 las, Antillas, y ha alcanzado a la parte norte de Centro America, M6xico,
y los Estados Unidos. Un tercer grupo (que incluye las tres algo extraviadas species
P. peramans, corculum, sanguinipes) origin6 en Sur America y ha alcanzado a las
Antillas Mayores y Centro America. Las species mas derivadas se encuentran en los
limits geograficos de cada grupos.

A new species of Protoneura Selys closely related to P. aurantiaca Selys has been
found in the foothills of the Talamanca range on the Caribbean coast of Costa Rica. The
new species has been taken well within the range of aurantiaca and only a few kilomet-
ers from an occurrence of that species. The proximity of occurrences of the two species
shows that the new species is not simply a geographic variant of aurantiaca. The new
species differs in having the brilliant orange of aurantiaca replaced by a bright sulfur
yellow which is more restricted on the head and on the abdomen, and by several struc-
tural details in both males and females.

Protoneura sulfurata, new species (Figs. 1, 2, 5, 7)

HOLOTYPE MALE. Head: dark iridescent green, with mandibles and center of ante-

Donnelly: New Costa Rican Damselfly


2 4

5 6

Figs. 1-7, Structural details and color pattern. Figs. 1 and 2, lateral and ventral
view of male terminal appendages of P. sulfurata. The ventral view has the right
superior appendages twisted slightly to show the interior tubercle in slightly different
views. Figs. 3 and 4, same for P. aurantiaca. Fig. 5, dorsal view of left side of female
mesothorax, showing mesostigmal lamina and prominence on mesepisternum. Fig. 6,
same for P. aurantiaca. Fig. 7, color pattern of male thorax of P. sulfurata.

clypeus brown; yellow as follows: distal rim of labrum; baso-lateral margins of ante-
clypeus, genae, front of frons bordering postclypeus and extending dorsally along edge
of eye to level of antennae; round triangular spots in front of antennae; ring at distal
end of scape.
Thorax (Fig. 7): prothorax black, boldly marked by yellow on middle lobes (except
for black mesal line between lobes) and hind lobe; pale ventral margin of proepimeron.
Pterothorax: mesepisternum yellow with black stripe on mid-dorsal carina extending
around posterior border; thin black line on posterior 2/5 along humeral suture, this
yellow extending forward to mesostigmal lamina and middle lobe of prothorax;
mesepimeron mainly black, yellow on dorsal 1/3, expanded posteriorly to cover most of
sclerite, black extending forward to form continuous band with mesinfraepisternum (ex-
cept for small yellow round spot on antero-dorsal corner) and proepimeron; metepister-
num pale in front, grading to yellow at rear; black on ventral 1/2 at rear thinning
forward and ending before level of metastigma; metinfraepisternum dark, grading to
pale ventrally; metepimeron pale; metasternum pale. Venation dark, pterostigma dark
red brown, rhombic, widened apically. CuP extending to just beyond cross vein descend-
ing from nodus; 3 antenodal spaces in proportion 2.1: 1: 1.6 proximall to distal). Legs:
fore femur yellow, middle femur pale with apical 1/2 yellow, hind femur pale with apical
1/3 yellow, with dark apices on femora and tibiae.
Abdomen: black, dorsal yellow line on 2, tiny dorso-basal yellow spot on 3, ventral
margins of terga pale, becoming distinctly yellow on apical 1/5 of 6 and all of 7 9;
terminal appendages (Figs. 1, 2) dark brown, superiors rounded in lateral view, tapering
to rear, shorter than inferiors, flattened laterally, with distinct meso-ventral part con-
taining black, highly sclerotized, sharply ridged mesal tubercle, the ridges curved in
mesal view; inferior appendage straight, tapering to rear, with subapical, mesally di-
rected triangular extension.

438 Florida Entomologist 72(3) September, 1989

ALLOTYPE FEMALE. Head: as in male; stripe on labrum broader, anteclypeus dull
Thorax: prothorax dark iridescent green, yellow ventral border of pronotum; hind
lobe erect, round, dark with central part of rim yellow. Pterothorax: dark, mesepister-
num with thin yellow-orange line above humeral suture extending posteriorly 1/2 length
of sclerite; mesepimeron with thin yellow line below humeral suture extending length
of sclerite except for anterior 1/6 and posterior 1/6, and with dorso-posterior round spot;
mesinfraepisternum dark; metathorax pale with dark dorsal and ventral borders of
metepisternum expanded and coalescing at rear; metinfraepisternum dark grading to
pale ventrally. Wings: as in male, antenodal spaces in proportion 2: 1: 1.5. Legs: pale,
fore femur with apical 2/5 dark, middle with dorsum of apical 1/2 dark, and hind with
dorsum of apical 1/6; apices of tibiae dark. Mesostigmal lamina (Fig. 5): rounded trian-
gular, low, laterally slightly inflated, antero-lateral rim slightly raised. Forward-di-
rected low, round prominence on anterior end of mesepisternum.
Abdomen: dorsum dark, sides of terga pale, conspicuously so on 1, 2, 8, and 9; lateral
round spot on 9; ovipositor reaching beyond end of segment 10.
Dimensions: Holotype male: abdomen 29 mm; hind wing 16.25 mm. Allotype female:
abdomen 25 mm; hind wing 17.25 mm.
VARIATIONS WITHIN TYPE SERIES. The type series of 10 males has abdomens ranging
from 26.5 to 31.25 mm and hind wings 15.5 to 17 mm. The series of 3 females has these
ranges 25 to 26 mm and 17.25 to 17.75 mm. The males vary slightly in extent of pale
color, with 6 males having the yellow on the front surface of the frons mesally divided
by a dark line up to 0.5 mm wide. Three males have the yellow on the ventral margin
of the 9th abdominal segment enlarged into a round spot. The 3 females show some
variation in the thoracic stripes, with 1 female having the stripe on mesepisternum
divided into anterior and posterior parts.
MATERIAL STUDIED. Holotype male, allotype female, 9 paratype males and 2 paratype
females: COSTA RICA: Lim6n Prov., Rio Pacuarito, 3 km. E. of Siquirres, 9-12 June
1986, Coll. T. and A. Donnelly. The locality was a tiny dammed tributary of the main
stream which was shallow and heavily vegetated. In addition to the Protoneura sul-
furata, there were several pairs of Psaironeura remissa, but few other odonates. The
holotype and allotype are deposited in the Florida State Collection of Arthropods,
ETYMOLOGY. The name, meaning "marked with sulfur yellow" is chosen so as to be
parallel in construction with the evry similar species aurantiaca.
DIAGNOSIS. The new species is very close to P. aurantiaca. The male is easily distin-
guished by its vivid sulfur-yellow color in contrast to the brilliant orange of aurantiaca;
the frons of sulfurata has the pale yellow confined to the front, whereas in aurantiaca
the pale orange extends dorsally to the top of the frons and 1/2 the distance to the
median ocellus; also the abdomen of aurantiaca has the 3rd segment vividly marked by
orange whereas in sulfurata this segment is almost entirely black. Females of sulfurata
differ from those of aurantiaca in thoracic color; the latter have the pale of the
mesothorax forming a broad orange line interrupted by a thin black line on the humeral
suture. There are minor structural differences. The pterostigma of sulfurata is widened
apically but not in aurantiaca. The male superior appendage of aurantiaca (Figs. 3, 4)
has an internal black tubercle with 2 or 3 sharply defined teeth, whereas in sulfurata
this structure extends further ventrally and is sharply ridged rather than toothed. the
mesostigmal lamina of aurantiaca (Fig. 6) is narrower than that of sulfurata and has
a high antero-mesal ridge; further the prominence on the cephalic end of the mesepister-
num is the end of an anteriorly divergent ridge and is more prominent than that of

Donnelly: New Costa Rican Damselfly


Calvert (1901-1908) provided the first study of the northern species of this genus
and was the author of five of the species. However, he discussed no interspecific re-
lationships, except for providing names based on a inscrutable metonomy which were
derived from the complexity of the female mesothorax and the supposed thoroughness
of the copulatory process. Williamson (1915) named a new species calverti which is close
to tennis and remains little known. He also split off three new genera from the originally
much broader generic definition. Further he noted on the heterogeneity of his newly
restricted genus and indicated implicitly that there were three groups of species. Cow-
ley (1941) discussed the difference between P. tennis and calverti. Donnelly (1961) de-
scribed P. ailsa but did not discuss its relationship with the known but still undescribed
P. viridis. Westfall (1964) described P. viridis and redescribed P. capillaris, noting
that, according to the character of the length of vein Cu1 (CuP in the Tillyard Fraser
notation used here), the two species can be differentiated. I have found this character
generally too variable for consideration in establishing relationships. In 1987 Westfall
described sanguinipes, but did not suggest its relationship with other species in the
genus. There exists to this date no discussion of species groupings among the entire
northern group of the genus.
The study of the new Protoneura raises questions concerning the relationships
among circum-Caribbean members of the genus. The species divide into two relatively
clear groups, with three aberrant species forming a possible third group. The first
group includes the South American to Mexican P. amatoria Calvert, sulfurata, the
Central American aurantiaca Selys, and the Central American to Mexican cupida Cal-
vert. This group is characterized by a rounded, laterally flattened male superior appen-
dage and a relatively small, straight, pointed inferior appendage with a lateral or subapi-
cal flat enlargement. The male superior appendage of each has a sclerotized internally
toothed tubercle, of which sulfurata has sharp ridges rather than proper teeth. Females
all have a raised prominence on the anterior ridge of the mesepisternum. In amatoria
this is a very prominent spine; in sulfurata it is least prominent. I would tentatively
conclude that amatoria stands near the base of this group and that loss of the mesepis-
ternal tubercle and reduction of the internal tubercle of the superior appendage are
derived characteristics.
The second group is far more diverse. The widespread South American (Brazil,
Venezuela, Guyana, Trinidad) P. tennis Selys is closely related to the rarer calverti
Williamson (Guyana; possibly reported in error from Trinidad by Geijskes, 1930). Both
have longer and straighter superior appendages with a ventral, more or less twisted,
less sclerotized keel. The inferior appendage is long and has a curved tip. I propose a
relationship between these two and the Lesser Antillean ailsa Donnelly, based on the
appearance of the inferior appendage, and on the elevated anterior part of the female
mesothorax. Within the Antilles, ailsa and the Greater Antillean viridis Westfall are
very closely related through the singular appearance of the elevated and swollen an-
terior part of the mesothorax. Females of these two are distinguished with difficulty
based only on their thoracic structures; I failed to notice (1961) the similarity of the
female thorax in these species. Both also share a relatively long male inferior appendage
with a slightly flared tip. However, their male superior appendages are substantially
different: in viridis the appendage is long and straight (like that of tenuis and calverti),
but in ailsa the appendage is shorter and rounded, somewhat like amatoria. I believe
the balance of characters, as well as distribution, places ailsa in this group in spite of
the singular male superior appendage.


440 Florida Entomologist 72(3) September, 1989

The Cuban P. capillaris Selys, a sister species of viridis, shares the salient struc-
tural characters of this group (Westfall, 1964). I would place P. cam Calvert, which
ranges from the United States to Central America, as a further, highly derived member
of this group. The male inferior appendage is shorter than other species in the group,
but the superior appendage is long and somewhat resembles that of viridis and capil-
laris. However, this appendage possesses a prominent baso-mesal spine unlike the
more mesal spine of those two species. I believe the most compelling reason to include
cara with this group is the swollen anterior part of the female mesothorax. If this
grouping can be supported, then the most likely origin of this group is South America
(a form close to tenuis) and the most derived species is cara; the distribution of the
group shows an advance through the Antilles and a subsequent invasion of Central
The three problematical species P. peramans Calvert (Central America), corculum
Calvert (M6xico and Central America), and sanguinipes Westfall (Hispaniola) provide
the greatest difficulty. I believe they are related more closely to each other than to any
other species. The male superior appendage of each is reduced and L-shaped, with a
relatively prominent baso-mesal swelling (almost a tubercle in peramans), and a prom-
inent sub-basal tubercle. The appendage and its prominences is smallest in corculum
and largest in sanguinipes. The male inferior appendage of each is relatively long. The
females of peramans and corculum share upwardly flared anterior rims of the mesostig-
mal laminae; in corculum these are folded backward. Both species have prominences
on the anterior margin of the mesothorax; in peramans these are spines, but they are
sharply in corculum. P. sanguinipes has a much simpler female thorax with prominent
but simple mesostigmal laminae. I place it tentatively as the most derived member of
this group because the male superior appendages are analogous to those of the other
two species. Another aberrant characteristic of this species is the relatively long CuP,
which extends an entire cell beyond the vein descending from the subnodus.
I do not know the diverse South American species sufficiently well to choose a
possible ancestral form for the third group, but the Peruvian paucinervis Selys shares
many of the characters of the male superior appendage. I believe this group is of
probable South American origin and has invaded Central America and, subsequently,
the Greater Antilles.
The above grouping and speculative derivation of the northern species of the genus
is based on subjective judgments as to which structural characters might be more
conservative than others. Clearly color or color pattern are unimportant, but it is less
clear whether the female thorax and the male superior appendage should be more
conservative than other structural characters. I have attempted to use both but suggest
that a biochemical investigation would be most interesting.


I am very grateful to Rosser Garrison, Minter Westfall, and Sid Dunkle for very
useful criticism of this paper.
Contribution No. 700, Bureau of Entomology, Division of Plant Industry, Florida
Dept. of Agriculture and Consumer Services, Gainesville, FL 32605.


CALVERT, P. P. 1901-1908. Neuroptera, Fam. Odonata, in Biologia Centrali
Americana, London, Porter & Dulati, 286 p.
COWLEY, J. 1941. A new species of Protoneura from Peru, and a review of the group
of Protoneura tenuis Selys (Odonata: Protoneuridae). Trans. Roy. Ent. Soc.
London 91(6): 145-173.

__ __ _____

I ___

Spencer & Havranek: New Venezuela Agromyzidae 441

DONNELLY, T. W. 1961. A new species of damselfly from St. Lucia, British West
Indies (Odonata: Protoneuridae). Florida Ent. 44(3): 119-121.
GEIJSKES, D. C. 1930. The dragonfly fauna of Trinidad in the British West Indies
(Odonata). Zool. Med. 14: 232-262.
WESTFALL, M. J. JR. 1964. A new damselfly from the West Indies (Odonata: Pro-
toneuridae). Quart. Jour. Florida Acad. Sci. 27(2): 111-119.
WESTFALL, M. J. JR. 1987. Protoneura sanguinipes spec. nov., a new protoneurid
damselfly from the Dominican Republic, West Indies (Zygoptera). Odonatologica
16(1): 93-97.
WILLIAMSON, E. B. 1915. Notes on Neotropical dragonflies, or Odonata. Proc.
U.S.N.M. 48: 610-638.

----- ---- -- - --- C i --- -- -*-


Exwell Farm, Bray Shop,
Callington PL17 8QJ,
Cornwall, England

U.N.E.T., Apartado 436, San Crist6bal, TAchira, Venezuela


The adult male and female of Melanagromyza amaranthi n. sp. and its puparia from
the State of TAchira, Venezuela are described. Besides the State of TAchira the fly has
also been collected in the States of Aragua and Merida, at an altitude range from 400
to 1,600 metres The host plant, Amaranthus dubious Mart. ex Thell., is common
throughout Venezuela and it has been collected at an altitude range from 120 to 1,800


El present trabajo describe los adults y la pupa de Melanagromyza amaranthi n.
sp., primero colectados para el Estado TAchira, Venezuela. Muesteos en los Estados
Aragua y Merida confirman su presencia tambien allf. Las muestras indican una dis-
tribuci6n altitudinal desde los 400 hasta los 1,600 m.s.n.m. La plant hospedera,
Amaranthus dubius Mart. ex Thell., se ha colectado desde los 120 hasta los 1,800
m.s.n.m. es una maleza comiin y con registros para casi toda Venezuela.

When studying insects attacking tomato on several farms in Venezuela, Daniela
Havranek also investigated the weed Amaranthus dubius Mart. ex Thell. (Havranek
1987). A. dubius is the most common species of 13 recorded for the genus Amaranthus
in Venezuela. Records of several herbaria (Universidad Central de Venezuela, Maracay,
State of Aragua; Universidad de Los Andes, Facultad de Ciencias Forestales, M4rida,
State of M4rida; and Herbario Nacional de Venezuela, Caracas, Distrito Federal), re-

Florida Entomologist 72(3)

vealed that it has been collected in almost all the States of Venezuela. The plant can
grow to a height of two metres, and it is found at altitudes from 120 to 1,800 metres.
It is a tropical cosmopolitan weedy species, but it is cultivated as a green vegetable in
West Africa and the Caribbean; in Java and other parts of Indonesia it is used as a
garden crop (BOSTID Reports 1984). One of the disadvantages of this plant is that it
is heavily attacked by many pests, especially defoliators. Puparia were found in the
stems of A. dubius and flies similar to Melanagromyza tomaterae Steyskal, 1972 were
reared from them. M. tomaterae is an agromyzid stem borer of tomatoes (Solanaceae).
It seemed improbable that a single species was involved in view of the high degree of
host specificity in the Agromyzidae. Detailed examination of the male genitalia of speci-
mens from the two hosts confirmed that the species on Amaranthus is an undescribed
new species. Both M. tomaterae and the new species belong to the large group of
greenish-black Melanagromyza which feed in the stems of a wide variety of hosts and
which can only be reliably identified by the examination of the male genitalia. In each
case the inside of the stems of the host are eaten by the larvae, thus causing death of
the plant. The new pest was first described using samples from San Crist6bal and
Rubio, State of TAchira, but also pupae have been found in El Cobre, State of TAchira
(23.V.1987, 1,300 metres above sea level); Maracay, State of Aragua (18.VII.1987, 400
metres above sea level), and in Bailadores, State of M6rida (22.IX.1987, 1,600 metres
above sea level). Bailadores was the only place where 30 pupae were collected in 10
stems, normally just one or two are collected in 10 stems or more. It is also important
to note that in each case this pest is competing with a lepidopteran pest in time and
space. The information so far obtained indicates that the new pest has a distribution
range from 400 to 1,600 metres. Some of the pupae were parasitised by Syntomopus
sp. (Hymenoptera: Pteromalidae) and two unidentified species of Hymenoptera

Melanagromyza amaranthi sp. n.
(Fig. 1)

Medium-sized, greenish black species, with head and wings typical of the genus.
Head.-Frons wider than eye, varying from 1.3 to 1.5 times eye width, only nar-
rowly projecting above eye in profile; 2 strong upper orbital bristles, 2 weaker lower
orbitals; orbital setulae conspicuous, in single row, reclinate; frontal triangle broad
behind, with apex at level of uppermost lower orbital; jowls relatively broad but vari-
able, from 1/5 to 1/7 vertical height of eye; third antennal segment small, round; arista
long, finely pubescent; eye in male with scattered pilosity at level of upper orbitals.
Wing.-Length from 2.5-2.75 mm in male to 2.75 mm in female; costa extending
strongly to vein M1 + 2; last section of M3 + 4 2/3 length of penultimate section; inner
cross-vein slightly beyond midpoint of discal cell.
Color.-Frons mat black, orbits and frontal triangle distinctly shining; mesonotum
greenish black, moderately shining, abdomen distinctly green; squamae and fringe sil-
very white, squamal margin scarcely differentiated; halteres black.
Male genitalia.-Aedeagus as in Fig. 1; epandrium with row of strong spines along
inner margin; surstyli broad, curving inwards, with numerous strong hairs or spines;
sperm pump large but with narrow blade.
Host.-Amaranthus dubius, larva feeding and pupating in stem; puparium whitish
yellow, posterior spiracular processes separated by almost twice their own diameter,
each with ellipse of about 14 large pores around short but strong central horn.
Holotype 6, Venezuela, State of TAchira, Rubio, Farm "La Tuquerena", 6. IX. 1986 (800
metres above sea level); paratypes: 1 6, same data; 1 Y San Crist6bal, Paramillo,
20.XI.1986 (900 metres above sea level); 1 9 State of Aragua, Maracay, 18.VII.1987;

September, 1989

Spencer & Havranek: New Venezuela Agromyzidae 443

aedeagus, ventral view. .

aedeagus, ventral view.

2 9, State of M6rida, Bailadores, 22.IX.1987, all ex stems of Amaranthus dubiu (D.
Havranek). Holotype and Y paratype in British Museum (Natural History), 1 C and 2
Y paratypes in K. A. Spencer's collection, 1 2 in collection of U.N.E.T.
Remarks.-This new species is of particular interest, as it is the first Melanagromyza
known with Amaranthus as host. A related species, M. chenopodii Spencer, 1963 was
described from Chenopodium ambrosioidea L. from Santiago, Chile which externally
resembles M. amaranthi (the Amaranthaceae and Chenopodiaceae are themselves
closely related) but the male genitalia differ considerably. When describing M.
chenopodii, the genitalia of a male from Ecuador were illustrated (Spencer 1963: Fig.
19) and it was clear that this was a further species which was not formally described at
the time. It now seems probable that this represents M. amaranthi.
With the relatively broad jowls, M. amaranthi runs in Spencer's (1973) key to
Venezuelan Melanagromyza species to M. tomaterae but M. consummate Spencer
from Bocon6, State of Trujillo also belongs correctly in the first alternative of the same
couplet (4). The male genitalia of these three species are quite distinct (Spencer 1973:
Figs. 17, 18: M. consummate and Figs. 49, 50: M. tomaterae).


I thank Dr. Boucek and Mr. Fergusson from the British Museum (Natural History)
for the Hymenoptera identification and to Mr. M. E. Bacchus for arranging it.

BOSTID Reports. 1984. Amaranth: Modern prospects for an ancient crop. National
Academy Press, Washington 80 p.

Florida Entomologist 72(3)

September, 1989

HAVRANEK, D. 1987. Melanagromyza tomaterae (Diptera: Agromyzidae) a tomato
pest in the States of TAchira and M6rida, Venezuela. Fl. Entomol. 70(2): 294-296.
SPENCER, K. A. 1963. A synopsis of the neotropical Agromyzidae (Diptera). Trans.
R. ent. Soc. London 115: 291-389.
SPENCER, K. A. 1973. The Agromyzidae (Diptera) of Venezuela. Rev. Fac. Agron.
(Maracay) 7(2): 5-107. (Venezuela).
STEYSKAL, G. C. 1972. Two new species of Melanagromyza Hendel (Diptera: Ag-
romyzidae) that bore in tomato stalks in Colombia and Ecuador. J. Washington
Acad. Sci. 62: 265-267.


'University of Florida, IFAS
Tropical Research and Education Center
18905 S.W. 280 Street, Homestead, Florida 33031

2Systematic Entomology Laboratory
Agricultural Research Service
USDA c/o U.S. National Museum NHB 168
Washington, D.C. 20560


Prodiplosis longifila Gagne (Diptera: Cecidomyiidae) is a polyphagous, Neotropical
species with a known range that extends into southern Florida. This gall midge feeds
on the flower ovaries of lime and can cause premature flower abscission. Details of its
biology on lime are given, the life stages illustrated, and the three larval instars de-
scribed in detail.


Prodiplosis longifila Gagne (Diptera: Cecidomyiidae) es una especie polifaga y neot-
ropical y su rango se extiende hasta el sur de la Florida. Esta mosquita destruye los
ovarios de las flores de los limones y puede causar caida premature de las flores. Se dan
detalles sobre su biologia en los limones, se hacen ilustraciones de los different estados,
y se described los tres estadios larvarios.

Prodiplosis longifila Gagn6 was recently reported feeding in flowers of lime, Citrus
aurantifolia (Christm.) Swingle (Rutaceae), in Dade and Collier Counties, Florida (Pefia
et al. 1987). Larvae completely destroy the ovaries of the flowers and may cause prema-
ture flower abscission. The only previous record of this species from the United States
was from wild cotton, Gossypium sp. (Malvacea), from Monroe Co., Florida (Gagne
1986). Prodiplosis longifila is otherwise known from Colombia and Peru, where it is
known to be a pest of tomatoes, potatoes, alfalfa, and other commodities (Gagne 1986).

Pefa et al.: A Gall Midge on Lime

In this report we present the first known details of the life history of P. longifila on
lime and describe the larval instars.


The biology of P. longifila was studied in a 0.7 ha, 10-yr old 'Tahiti' lime orchard in
Homestead, Florida and in the laboratory at the Tropical Research and Education
Center, Homestead, Florida. Approximately 30 trees were used in this study. Observa-
tions were made over a period of 86 days (November 1987 through January 1988) for
five generations of the gall midge.
Lime flowers in their first stages of development (less than 1 mm in diameter) were
bagged with nylon mesh. Flowers were left exposed for 12 h and egg masses collected
after females oviposited. Egg masses (n = 14) were transferred to the laboratory and
eggs n = 258) placed individually in petri dishes (5.2 cm in diameter) lined with filter
paper. Petri dishes were held at 27C and 842 RH and observed daily. After eclosion
each larva was removed and transferred to similar petri dishes with excised flowers.
Larvae were removed two times during the day to determine the duration of each
instar. Full-grown larvae from a set of infested flowers were allowed to drop into one
of 12 360 ml carton containers, each containing 400 g of previously sterilized 'Rockdale'
soil. Pupae were collected from the soil surface and from different soil depths (1.5-2.9
cm, 3.0-5.4 cm, and 5.5-8.0 cm). Adult emergence was determined in the field by placing
infested flowers on plastic containers (43 x 59 x 26 cm) filled with 'Rockdale' soil.
Clear-plastic circular plates (24 cm diam.) coated with Tanglefoot were placed on top
of the containers. Adults trapped on the plastic plates were counted every 2 h during
a 24 h period on five separate days (November 19, 20, 21, 1987 and January 14, 15,
1988). Adults kept in the laboratory were maintained in a rearing chamber at 270C and
842 RH with ca. 12 h photoperiod. Each adult was kept in a petri dish lined with filter
paper and provided with ca. 0.01 ml of honey and 0.01 ml water or left without food.
Larvae used for detailed descriptive study were mounted on glass slides in Hoyer's
mounting medium or Canada balsam for study. Measurements were made of 20 speci-
mens of each instar. Representative specimens have been deposited in the National
Museum of Natural History, Washington, D.C.


Eggs are transparent, elongate-ovoid, 0.265 0.035 SE mm x 0.096 0.002 SE mm
(Fig. 2). They are deposited on stamens or styles, usually of flowers that are 0.46-0.65
cm in diameter [x2.05[8] ; P<0.05] or in flowers in which a small opening in the corolla
allows the style to protrude. One to 59 eggs (mean = 12.62 SE) were found per
infested flower under laboratory conditions. Larvae hatched in 1.40.045 SE days.
Larvae are almost transparent when newly closed, turn white 1.2 days later and
become yellowish to orange when full-grown (Fig. lc). There are three instars. Major
anatomical features common to all three (Figs. 5-13) are a small head capsule with
one-segmented, conical antennae, a neck segment, three thoracic segments, and nine
abdominal segments, each segment with papillae characteristic for the Cecidomyiini in
number and position (Gagn6 in press). The spiracles of the eighth abdominal segment
are situated posteriorly.
The first instar (Figs. 5-7) ranges from 0.40-0.92 mm in length. The head capsule is
0.045 0.003 mm wide at the posterior end. This instar has only one pair of spiracles,
on the eighth abdominal segment (Fig. 5), and that is the most conspicuous difference
between this instar and the remaining two.
The second instar (Figs. 8-10) ranges from 0.76-1.85 mm in length. The head capsule

Florida Entomologist 72(3)

Fig. 1-4. Prodiplosis longifila. 1, adult. 2, eggs. 3, third instar. 4, pupa.

is 0.0500.005 mm wide at its posterior end. This instar has the full complement of
spiracles for cecidomyiid larvae, one pair on the first thoracic segment and one pair on
each of the first through eighth abdominal segments. It differs from the third instar in
lacking a spatula and having the corniform pair of papillae on the terminal segment less
strongly developed relative to the other three pairs.

September, 1989


Pefa et al.: A Gall Midge on Lime

Figs. 5-10. Larvae, Prodiplosis longifila. 5-7, First instar: 5, dorsal view; 6, head
through second thoracic segment, ventral; 7, abdominal segments seven through nine,
dorsal. 8-10, Second instar: 8, dorsolateral view; 9, head through second thoracic seg-
ment, ventrolateral; 10, abdominal segments seven through nine, dorsolateral. Line =
0.1 mm.

The third instar (Figs. 11-13) ranges from 1.15-1.90 mm in length. The head capsule
is 0.0500.005 mm wide at its posterior end. The spiracular system is similar to that
of the second instar. The distinguishing feature of this instar is the clove-shaped spatula
on the venter of the first thoracic segment (Fig. 12). The papillar setae, except for those


448 Florida Entomologist 72(3) September, 1989

o o

o o

11 .



Figs. 11-13. Third instar, Prodiplosis longifila. 11, dorsal view with the ventral
spatula dotted in to show relative size and position; 12, head through first thoracic
segment, ventral; 13, abdominal segments seven through nine, dorsal. Line = 0.1 mm.

of the terminal segment, are shorter relative to the width of the papillar bases than in
previous instars.
The head capsule width of cecidiomyiid larvae usually becomes significantly larger
in each instar (Gagne and Hatchett in press, Roskam 1977, 1979, Wilson 1966), but in
P. longifila the head of the second instar is as wide as that of the third. First instar
head capsules are sometimes also as wide as those in following instars. Antennae are
similarly shaped in each instar.
Two days after hatching, 17 of 20 larvae collected were first instar, three were
second instars. Three days after hatching, 12 of 20 larvae were second instars and eight
were third instars. Four days after hatching all larvae were third instars. The entire
larval stage last 91.63 days.
Larvae feed on the surface of the ovary, pistils, and stamens of lime flowers. One
to 66 larvae were found per infested flower (24.264.15 SE). Mature larvae drop to the
ground, where they penetrate the soil more frequently at a depth of 1.5 cm than at


Pena et al.: A Gall Midge on Lime





c 2.0-


0. 1.0-





i 0


Time of Day

Fig. 14. Average daily catches of Prodiplosis longifila adults every two h.

other depths (x2.005[3] = 7.81; P<0.005). They then spin a white cocoon, which some-
times incorporates sand grains.
Pupae are 0.85-1.00 mm long and pale yellow when newly molted (Fig. 4). The head
and thorax turn black 3.020.92 SE days later. This stage lasted 4.111.22 SE days
unless parasitized.
Adults are about 1.5 mm in length (Fig. 1). Wing length is 1.420.04 mm in males,
1.530.02 mm in females. A technical description of this stage is given in Gagn6 (1986).
Peak emergence occurred between 1700 and 2300 hours, but a few adults emerged at
other times (Fig. 14). Temperatures recorded during periods of adult activity ranged
from 17- 200C. Relative humidity ranged between 69-98%. The female to male sex ratio
of emerging adults fluctuated between 70:30 and 50:50 (n = 87-40). Adults reared in
the laboratory survived 1.060.24 days if not fed; if held in vials with available honey
and water, they survived 8.301.21 days, with a maximum of 16 days.
Prodiplosis longifila is parasitized by Synopeas sp. (Platygasteridae), an egg-larval
parasitoid. Parasites emerged 14-16 days after pupation of the gall midge.


We thank Deborah Leather Roney, Arlington, Virginia, for the detailed larval draw-


450 Florida Entomologist 72(3) September, 1989

ings and L. Stange, Florida Department of Agriculture and Consumer Services, Divi-
sion of Plant Industry, Gainesville, for identifying the parasitoid of P. longifila. We are
grateful to R. M. Baranowski, University of Florida, Homestead, P. E. Boldt, Agricul-
tural Research Service, USDA, Temple, Texas, N. E. Woodley, Agricultural Research
Service, USDA, Washington, D.C., H. C. Roskam, University of Leiden, The Nether-
lands and L. F. Wilson, East Lansing, Michigan for commenting on a draft of the ms.
The work was supported by a grant from the Florida Lime and Avocado Committee.
This paper is Florida Agricultural Experiment Station Journal Series No. 9573.


GAGNE, R. J. 1986. Revision of Prodiplosis (Diptera: Cecidomyiidae) with descrip-
tions of three new species. Ann. Entomol. Soc. Am. 79: 235-245.
- In press. The plant feeding gall midges of North America. Cornell University
Press, Ithaca, New York.
GAGNI, R. J. AND J. H. HATCHETT. In press. Larval instars of the Hessian fly (Dipt-
era: Cecidomyiidae). Ann. Entomol. Soc. Am.
PENA, J. E., R. M. BARANOWSKI, AND R. T. MCMILLAN, JR. 1987. Prodiplosis
longifila (Diptera: Cecidomyiidae) a new pest of citrus in Florida. Florida En-
tomol. 70: 527-529.
ROSKAM, H. C. 1977. Biosystematics of insects living in female birch catkins. I. Gall
midges of the genus Semudobia Kieffer (Diptera: Cecidomyiidae). Tijdschr. En-
tomol. 120: 153-197.
1979. Biosystemtics of insects living in female birch catkins. II. Inquiline and
predaceous gall midges belonging to various genera. Netherlands J. Zool. 29:
WILSON, L. F. 1966. Life history, habits, and damage of the boxelder leaf gall midge,
Contarinia negundifolia Felt (Diptera: Cecidomyiidae) in Michigan. Canadian
Entomol. 98: 777-784.


Florida State Collection of Arthropods,
Division of Plant Industry,
P. O. Box 1269
Gainesville, Florida 32602


The genus Dinmares Hagen is divided into two genera, Dimares with one species in
Brazil and Argentina, and a new genus Millerleon with three species in the coastal
desert of Peru. A world-wide key for the genera in the tribe Dimarini is presented based
on adults and larvae. Also, a key for the species in the genus Millerleon is presented.
A diagnosis of the tribe, the two American genera and the American species are pro-
vided with new records for the species. Data are provided about the biology and mor-
phology of the larvae of Dimares elegans and Millerleon bellulus.

S- r, JT i . :"; *1|

- *



Stange: New World Dimarini 451


Se divide el g6nero Dimares Hagen en dos g6neros, Dimares que consta de una
especie en Brasil y Argentina, y Millerleon, un genero nuevo con tres species en el
desierto costal del Perd. Se present una clave para los generos del mundo de la tribu
Dimarini basada sobre los adults y las larvas. Tambi6n, se present una clave para las
species de Millerleon. Un diagnosis de la tribu, los dos g6neros Americanos y las
species Americanas son provistas con registros nuevos de las species. Se proporcion6
datos sobre la biologia y morfologia de las larvas de Dimares elegans y Millerleon

In 1982 Miller and Stange discovered Dimares larvae in the Peruvian Coastal Desert
and reared two species. These larvae, when compared to larvae of Dimares elegans
(Perty) from Argentina, possess so many odd characteristics that it is apparent that the
two groups of Dimares are not congeneric. The adults also show significant differences
in the form of the male paramere, presence or absence of the pilula axillaris, and in the
development of the pretarsal claws. Accordingly, a new genus is proposed for the Peru-
vian coastal desert species. A key to genera, based on adults and larvae, is provided as
well as a key to the species.

Tribe Dimarini Navas

Dimarini Navas. 1914. Mem. R. Acad. Barcelona 11:107
Echthromyrmicini Markl, 1954. Verh. Naturf. Ges. Basel 65:217. New synonymy.
Echthromyrmicinae Holzel, 1972. Beitr. naturk. Forsch. SudwDtl. (Suppl.) 1972 (1):8.
Further Description: Markl 1954:217-218.
Diagnosis: ADULTS: Pronotum wider than long; legs stocky, femoral sense hair
absent; tibial spurs well developed; tibial spurs and pretarsal claws thick, well de-
veloped; pretarsal claws strongly curved, not capable of closing upon tarsus; labial
palpus elongate, distal palpomere thread-like, sensory area often slit-like; male abdomen
with non-eversible glandular openings on pleura between segments 5 & 6, 6 & 7, and 7
& 8, without associated setae; male ectoproct without postventral lobe; female posterior
gonapophysis less than 1.5X longer than wide; female ectoproct and lateral
gonapophysis with strong digging setae; forewing vein CuP fuses with vein IA a short
distance from wing base; hindwing vein CuP curves anteriorly to fuse with posterior
fork of MP + CuA a short distance after fork; hindwing radial sector arises near base,
1-3 presectoral crossveins.
LARVAE: Labial palpus shorter than basal width of mandible; antenna short, flagello-
meres all wider than long,, mandible with 2 or 3 teeth,, sternite IX with or without
highly modified digging setae.
Discussion: This tribe has 3 known genera, Echthromyrmex from the Ethiopian and
Oriental Regions and two genera from the New World. Larvae are known only for the
New World genera. The tribe belongs to the subfamily Palparinae. The combination of
the absence of the femoral sense hair (except Maulini), condition of the hindwing vein
CuP, and thread-like labial palpus (not thread-like in some Palparini) are diagnostic
characteristics of the Palparinae of which only the tribe Dimarini is represented in the
New World. It differs from the largest tribe of the subfamily, Palparini, in the fusing
of forewing vein 1A with MP + CuP. However, one species in the Palparini, Pal-
panrtius concinnus Peringuey from southern Africa, has wing venation as in the Dima-
rini. A third tribe in the subfamily, Maulini, is similar to the Dimarini in wing venation

Florida Entomologist 72(3)

except in Maulini the origin of the radial sector of the hindwing is much more distad
from the base. The genera can be identified with the following key.



1. Sensory opening of distal palpomere of labium does not reach apex of pal-
pomere; Old World ............................................ Echthromyrmex McLachlan
Sensory opening of distal palpomere of labium extends around apex and
nearly reaches opposite side; New World .............................................. 2
2. Pretarsal claws of hindleg much longer than hind basitarsus; (Fig. 2); male
paramere with long hook, toothed apically (Fig.4); no teeth along mesal
margin; male pilula axillaris absent; sexual dichromatism present, males with
essentially unmarked wings, females usually with numerous brown spots or
bands; Brazil south to Argentina ........................................ Dimares Hagen
Pretarsal claws of hindleg shorter than hind basitarsus (Fig. 1); male para-
mere without hook, 12 or more teeth along mesal margin (Fig. 5); male pilula
axillaris well developed; both male and female with pigmented wing spots or
bands; Peruvian coastal Desert (southern Ecuador to northern Chile) ..........
..................................................................................... M illerleon Stange


Mandible with 3 teeth (Fig. 15); sternite VIII with well developed submedian
tooth; sternite IX with 2 pairs of highly modified digging setae (Fig. 15);
Brazil to Argentina ............................................................. Dimares Hagen
Mandible with 2 teeth (Fig. 13); sternite VIII without submedian tooth; sternite
IX without highly modified digging setae (Fig. 12); Peruvian Coastal Desert .
.................................................................................. M illerleon Stange

Dimares Hagen 1866

1866. Dimares Hagen. Stettin. ent. Ztg. 27:372, Type species: Myrmeleon elegans
Perty, by original designation.
1909. Banks. J. New York Ent. Soc. 17:1-2 (Key to species)
1913. Navas. Mis. Arc Meridien Equatorial Amer. Sud 10(1):71 (Key to species)
1914. Navas. Mem. R. Acad. Barcelona 11:101 (in tribe Dimarini)
1976. Riek. Journal Australian entomological Society 15:297 (in Dendroleontinae)
Diagnosis: Adults: Sexual dichromatism present, males without pigmented wing
bands, females usually with pigmented wing bands; frons with several scattered setae;
distal labial palpomere with slit-like opening extending around apex to opposite side;
basitarsus of foreleg about 2.0X longer than middle diameter, that of hindleg about 3.OX
longer than middle diameter of tarsus; hind tibial spurs as long as or longer than basitar-
sus; pretarsal claws at least 1.5X longer than hind basitarsus; male pilula axillaris
absent; male paramere with long hook, 2-4 teeth apically. Female terminalia with pre-
genital plate membranous; gonapophyseal plate long and slender. LARVAE: mandible
with 3 teeth; ventral head capsule with numerous setae; sternite VIII with submedian
tooth; sternite IX with 2 pairs of highly modified digging setae posteriorly (Fig. 15).
Systematics: This genus appears to have only one geographically variable species,
Dimares elegans (Perty). This genus is unusual because the males and females show
marked sexual dichromatism, the males having unmarked wings and most of the females


September, 1989

Stange: New World Dimarini

have extensive brown markings. This led to a duplicity of names. Walker (1859) de-
scribed each sex as a different species. Subsequent workers, especially Longinos Navas
and Nathan Banks, were also unaware of this sexual dichromatism.

Dimares elegans (Perty)
(Figs. 2, 4-6, 15)

1833. Myrmeleon elegans Perty, Delect. animal. articul. Brasil 3:125. Holotype female,
Brasilia (Munich!).
1853. Walker. List neuropterous insects British Museum p. 395 (repeat of Latin de-
1859. Myrmeleon conicollis Walker Trans. ent. Soc. Lond. 5:188. Holotype female,
Santarem (Brazil) (BMNH!).
Myrmeleon albidilinea Walker, Trans. ent. Soc. London 5:189. Holotype male,
Amaz. (BMNH!).
1861. Myrmeleon congruus Hagen, 1861. Smith. Misc. Coll. 4(1):326.
1866. Hagen, Stettin, ent. Ztg. 27:372 (in genus Dimares), 437 (congruus, nomen
1867. McLachlan, J. Linn. Soc. (Zool.) 9:281 (conicollis = elegans).
1909. Banks, J. New York ent. Soc. 17:146 (in Key to species).
1912. Dimares elegans lepida Navas. Broteria 10:41 Fig. 3. Syntypes, Catamarca &
Mendoza, Argentina (Copenhagen).
1914. Dimares erythrostigma Navas, Broteria 12:47, Fig. 2. Holotype female, Solidade,
Brazil (Vienna!).
1915. Navas. Mem. R. Acad. Barcelona (3)12:125 (lepidus= valid species)
1915. Navas. Rev. Acad. Madrid 17:293 (Records from Argentina)
1920. Dimares hageni Banks, Bull. Mus. Comp. Zool. Harv. 64:330.
Syntypes, Chapada, Brazil, H. H. Smith (MCZ). (lepidus = elegans).
1920. Esben-Petersen, Bull. Ann. Soc. R. ent. Belgique 60:190 (erythrostigma=
1920. Navas. Ann. Soc. cient. argent. 90:16, 57 (Records from Argentina).
1920. Navas, Estudios 22:359 (Records from Argentina).
1922. Navas, Rev. Acad. Madrid 19:255, 256 (Records from Argentina).
1922. Navas, Arx. Inst. Cienc. 7:182 (Records from Argentina).
1923. Navas, Rev. Mus. Paulista 13:768-771) (Taxonomy: hageni= elegans).
1926. Navas, Estudios 32:105 (Records from Argentina).
1928. Navas, Estudios 35:140 (Records from Argentina).
1930. Navas Rev. Soc. ent. argent. 3:127 (Records from Argentina).
1954. Markl, Verh. naturf. Ges. Basel 65:196, 203, 204, 217, Figs. 10, 41, 44, 65 (wing
1967. Stange, Acta Zool. Lilloana 22:43 (albidilinea= elegans).
1976. Riek. Australian J. Zoology 15:344, Fig. 5 notaa, wing base).
1976. Stange et al. Acta Zool. Lilloana 32:109 (Larva).
1984. Stange, Nat. Geographic Soc. Res. Reports 17:69 (Record from Argentina).
Diagnosis: Length of body 30-40 mm; forewing length 20-36 mm, greatest width 7-10
mm. Face yellowish brown with darker brown band below, laterad and above antennal
fossae, darkest mesad of fossae, sometimes dark brown on labrum; vertex with anterior
vertex row of three dark brown scar marks, middle mark extends posteriorly to middle
row of vertex markings which consist of five irregular markings; posterior row with
median dark spot, sometimes submedial dark spot; antenna nearly all dark brown except
scape and pedicel apically; labial palpus yellowish brown mesally, dark brown externally;
nota yellowish brown with dark brown stripe medially and sublaterally; legs pale brown


Florida Entomologist 72(3)

rr .4

1 2

Fig. 1. Hind tarsus of Millerleon subdolus; Fig. 2. Hind tarsus of Dimares elegans;
Fig. 3. Female terminalia (exploded ventral view) of Millerleon subdolus; Fig. 4. Male
genitalia of Dimares elegans; Fig. 5. Male genitalia of Millerleon subdolus.

except subbasal dark spot on exterior face of tibia and coxae which are dark basally and
posteriorly, with dark stripe on lateral face; abdominal sclerites nearly completely dark
brown except pale brown on terminalia; pretarsal claws and tibial spurs reddish brown;

September, 1989

Stange: New World Dimarini



Figs. 6-10. Wings of Dimarini. 6. Dimares elegans (male); 7. Dimares elegans
(female); 8. Millerleon subdolus; 9. Millerleon bellulus; 10. Millerleon pretiosus.



Florida Entomologist 72(3)

wing veins and crossveins nearly all dark brown (males) or pale in non-pigmented areas
(females), stigma of males white, sometimes pinkish (Brazil); pronotum with erect black
setae on margins; mesoscutum with numerous erect bristles; scutelli with elongate white
setae posteriorly; legs with black setae shorter than width of segment at point of origin
except a few on hindtibia; femora with longer black setae mostly on closing face, tibiae
with black setae on both closing and exterior faces; abdomen with longest setae on
tergites I-II and terminalia, elsewhere very short; female ectoproct and lateral
gonapophysis with well developed digging setae; male ectoproct with long setae post-
eriorly. Greatest ocular width much shorter than interocular distance; labial palpus very
long, penultimate palpomere longer than distal one, about as long as greatest head
width (including eyes); antenna with about 28 flagellomeres, flagellomeres 1-8 longer
than wide; pronotum about 2X wider than long; legs about equal length; tibial spurs of
hindleg somewhat shorter than basitarsus (Argentina) or longer than basitarsus (Brazil);
pretarsal claws shorter than fore basitarsus (Argentina) or almost as long as distal
tarsomere (Pernambuco); hindwing about as long as forewing, in repose apex of hindw-
ing extends beyond that of forewing; female usually with extensive brownish suffusion
in form of short bands or spots; male wing always without suffusion; male abdomen
somewhat longer than wings, without tufted pore plates; female abdomen somewhat
shorter than wings; male sternite IX transverse, emarginate medially; male genitalia
(Fig. 4), with paramere with 3-4 apical teeth; female terminalia with posterior
gonapophysis about as long as wide.
ARGENTINA: Catamarca: 6 km S Santa Maria, III.21.1974, L. Stange (2 male, 5 female,
3 larvae-SC); Belen XII.19.1971, C. Porter & L. Stange (2f-SC, IML). Tinogasta,
II.8.1966, L. Stange (6m, 2f-SC, IML); Cordoba: Dique Los Molinos, 1.10.1975, Willink
(If-IML); Sierra, 1.10.1927, Williner (If- SALTA); Villa Dolores, II.7.1965, Kohler (2m-
IML); Entre Rios: Concordia, Salto Grande, 1.1975 (lm-SC); La Pampa: Sierra Lihuel
Calel, II.1.1968, L. Stange (2m, If-SC); La Rioja: Campanas, II.1966, L. Stange (1m-
SC), Capital, III.6.1970, C. Porter & L. Stange (lm-SC); Mazan,III.14.1969, A. Teran
& L. Stange (lm-SC); Mendoza: Pareditas (Ruta 40), 1.18.1975, A. Willink (lf-IML);
Rio Negro: Chimpa, 1.16.1975, A. Willink (1f-IML); Salta: Las Mesitas, II.17.1962, A.
Willink (Im-IML), Santiago del Estero: Las Termas, 1.12.1966, L. Stange (lm-SC); Los
Tigres, 1.16.1970, R. Golbach (3m-SC, IML), San Luis: Lujan, II.23.1966, L. Stange
(Im-SC); San Juan: Chicuma, 1.20.1959 (If-SC); Tucuman: Vipos, III.21.1971, P.
Fidalgo (Im-SC); Trancas, (2f-La Plata); BOLIVIA: Santa Cruz: El Palmar Oratoria,
1.25.1980, L. Stange (1m, 5f-SC)
BRAZIL: Ceara: Barbalha, V.1960, M. Alvarenga (2m, 6f-SC); Goias: S. Isabel do
Morro, Iha do Bananal, VI.1961, M. Alvarenga (2m, If-SC); 48 km., 124 km. S. Peixe,
VI.2.1956, F. Truxal (3m, 2f-LACM, SC); Mato Grosso: Barracao Queimado, XI.1960,
M. Alvarenga (3m, If-SC); Gustavo Dutra, Cuiaba, XI.1963, M. Alvarenga (3f-SC);
Pernambuco: Petrolandia, V.1969, M. Alvarenga (1m, 3f-SC); Rondonia: Vilhena,
XI.1960, M. Alvarenga (2m-SC).
URUGUAY: Arenitas Blancas, 11.3.1962, P. San Martin (If-SC).
Discussion: This species shows geographic variation in its extensive range from
Pernambuco, Brazil to Rio Negro, Argentina. The populations occurring in the Suban-
dean Desert and Chaco of Argentina have somewhat less extensive wing markings in
the female than most of the populations occurring in Brazil. Also, the stigma is white

Fig. 11-14. Third instar larva of Millerleon bellulus. 11. Dorsal view; 12 Ventral
view of abdominal apex; 13. Ventral view of head; 14. Ventral view.
Fig. 15. Third instar larva of Dimares elegans (Ventral view).

September, 1989


Stange: New World Dimarini





,- -^A"^-.- -
'SR Ifr~
t.U-ts 4-* ;
w-.'rft^ 1I<*' 7
-^q y


qL~ a



458 Florida Entomologist 72(3) September, 1989

in the Argentina males often somewhat reddish in the Brazilian populations. Specimens
seen from Pernambuco, Brazil, appear to have less extensive wing markings in the
females. About 10% of the females have the wing markings much reduced or even
absent. The tibial spurs are shorter than the hind basitarsus in the Argentine populations
whereas in Bolivia and Brazil they are longer than the hind basitarsus. This may be
clinal variation since the northernmost populations from Pernambuco appear to have
the best developed tibial spurs and pretarsal claws. The male genitalia are very similar
among all of these populations, with slight differences in the length and width of the
paramere hook.
Biology: Larvae (Fig. 15) of Dimares elegans were found in fairly deep sand between
sand dunes in Argentina. The larvae move slowly, both backwards and forwards. They
were not reared but their structural characteristics identify them with a high probability
of accuracy.

Millerleon Stange, new genus
(Figs. 1, 3, 5, 8-14)

Type-species: Myrmeleon subdolus Walker, by present designation.
Description: ADULTS: Sexual dichromatism absent males and females with equivalent
wing markings; frons without setae; distal labial palpomere with slit-like opening ex-
tending around apex to opposite side; basitarsus of foreleg at least 3.OX longer than
middle diameter of tarsus; hind tibial spurs shorter than basitarsus; pretarsal claws no
longer than hind basitarsus; male pilula axillaris present; base of hindwing posterior
vein and male pilula axillaris with elongate hair-like setae; male paramere without hook,
12 or more teeth along mesal margin of paramere; female terminalia with small
sclerotized pregenital plate (Fig. 3); gonapophyseal plate absent. LARVA: mandible
with 2 teeth; ventral head capsule nearly glabrous; sternite VIII without submedian
tooth; sternite IX without highly modified digging setae posteriorly.
Discussion: There appear to be 3 described species. The species are very similar
structurally with identical male genitalia. There are differences in the wing shape, and
to a lesser degree, the wing markings which vary considerably. Tegumental characters
of the male abdomen appear to be important differences between species. More speci-
mens and study are needed to clarify the species definitions in this genus. All the species
are restricted to the Peruvian coastal desert (south Ecuador to northern Chile). This
genus is named for Robert Bruce Miller in recognition of his outstanding field work with
larval and adult antlions.

Key to Species of Millerleon Stange

1. Hindwing less than 3.5X longer than greatest width which occurs about mid-
point of wing (Fig. 8); apical one-fifth of wing mostly with solid dark brown
bands, most of hypostigmatic cell completely dark brown suffused; male
abdomen with most tergites and sternites (especially toward posterior end)
with numerous pore plates, mainly associated with setal bases, giving ab-
domen a scaly appearance; south Ecuador to central Peru ...... subdolus (Walker)
Hindwing more than 4.OX longer than greatest width which occurs well
beyond midpoint of wing (Figs. 9, 10); apical one-fifth of wing mostly pale
brown with numerous dark brown suffused areas, hypostigmatic cell mostly
not suffused; male abdomen without tufted pore plates at least beyond tergite
III usually smooth toward posterior end ............................................. 2

Stange: New World Dimarini 459

2. Male pilula axillaris with head bearing setal mat about 3.OX wider than base
of pedicel; forefemur dark brown; south Peru to north Chile ... pretiosus (Banks)
Male pilula axillaris with head about 2.OX wider than base of pedicel; fore-
femur light brown; south Ecuador to central Peru ................ bellulus (Banks)

Millerleon bellulus (Banks) 1908, New Combination
(Figs. 9, 11-14)

1908. Dimares bellulus Banks Proc. ent. Soc. Wash. 9:30. Holotype, Posorja, Ecuador
1913. Dimares amoenus Navas, Mis. Arc Mer. Equatorial Amer. Sud 10(1):70. Syn-
types: If, Ecuador, Staudinger & Rolle (Barcelona) 2m, Perou, Paita et tablazo
de Paita 50-80m, P. Rivet, 1906 (Paris). New synonymy.
Diagnosis: Forefemur light brown; hindwing about 4.OX longer than greatest width,
which occurs well beyond midpoint of wing; apical one-fifth of wing mostly pale brown
with numerous dark brown suffused areas; hypostigmatic cell mostly not suffused; male
pilula axillaris with head about 2.OX wider than base of pedicel; abdomen dull with pale
spicules on most sternites and tergites, dark brown tufted pore plates only on tergites
and sternites I-III.
Material examined: ECUADOR. Guayas: 27 mi. N. Santa Elena, 1.30.1955, E.
Schlinger & E. Ross (2m-CAS, SC); Posorja, III, Campos (Im-SC), PERU. Piura:
Puerto Pizarro, 14 km. N. Tumbes, V.6.1959, W. Weyrauch (4m 5f- IML,SC); Lam-
bayeque: 14 km. N. Mocupe VII.1985, R. B. Miller & L. A. Stange (2 reared females-4
larvae-SC,MC), La Libertad, Pacasmayo, V.6.1959, W. Weyrauch (If-SC).
Biology: Larvae (Fig. 11) were found about 18 inches under loose sand that accumu-
lated in animal burrows. These special holes, usually found around small bushes or bases
of small sand dunes, have a lot of organic debris (mostly leaves) and a variety of beetle
larvae. Larvae feed underground. A photograph of one of the habitats is published in
Stange (1984, Fig. 2). These larvae are very sluggish, but move forwards and back-
wards. Only 2 larvae were reared after 3 years in the laboratory. The time period in
the cocoon was 14 or 33 days. The adult lived 8 days after emerging without benefit of

Millerleon pretiosus (Banks) New Combination
(Fig. 10)

1908. Dimares pretiosus Banks. Journal New York Ent. Soc. 17:1. Holotype, Mollendo,
Peru, December (MCZ).
Diagnosis: Forefemur dark brown; hindwing about 4.0X longer than greatest width
which occurs well beyond midpoint of wing; apical one-fifth of wing mostly pale brown
with numerous dark brown suffused areas; hypostigmatic cell mostly not suffused; male
pilula axillaris with head about 3.0X wider than base of pedicel., abdominal tergites I-II
dull brown with pale spicules; tergites III-VII mostly shiny brown.
Material examined: CHILE. Tarapaca: Cuya, Quebrada de Camarones, Arica
II.25.1976, N. Hitchins (1 male-FSCA); km 12, Valle de Azapa, Arica, III.6-23.1977,
N. Hitchins (lm-FSCA); Azapa Grande, II.7.1964 (1m,' If-FSCA); Azapa, 11.20,1948
Systematics: The abdominal tegumental sculpture is different from M. subdolus and
from M. bellulus. The longer wings and other details point to a closer association with
M bellulus but the larger male pilula axillaris is a notable difference from that species.

Florida Entomologist 72(3)

Millerleon subdolus (Walker) New Combination
(Figs. 1, 3, 5, 8)

Myrmeleon subdolus Walker 1853. List Neuropterous insects British Museum p. 395.
Holotype, Lima, Walker Coll. (BM!)
Dimares formosus Banks 1908. Proc. ent. Soc. Washington 9:31. New synonym.
Holotype, Posorja, Ecuador, Campos (MCZ).
Dimares venustus Banks 1908. Proc. ent. Soc. Washington 9:31. New synonym.
Holotype, Posorja, Ecuador, Campos (MCZ).
Dimares nummatus Navas 1912. Ann. Soc. Scient. Bruxelles 36:229, Fig. 16 (hindwing)
(after Stange 1969:189). Holotype male, S. Elena, Ecuador (not Galapagos!), 1,
4.II.1876 (Vienna!)
Dimares decorus Navas 1913. Mission geographique Acad. Sci. 10:69, P1. iv, Fig. 6
(hindwing). New synonym. Holotype, Paita et Tablazo de Parita, Peru, 50-80 m,
1906, P. Rivet (Paris)
Dimares formosus punana Navas 1933. Acad. Cienc. Nat. Madrid 29:196 Fig. 15
(hindwing). New synonym. Syntypes, Puna Vieja, Ecuador, Campos (Barcelona)
Taxonomy: Hagen 1860:360 (in Palpares); 1866:403 (in Dimares; McLachlan 1867:281
(subdolus not= elegans); Esben-Petersen 1920:491 (nummatus= formosus)
Distribution: Ecuador (Campos 1922:72; Navas 1935:361).
Diagnosis: Forefemur yellowish brown; hindwing less than 3.5X longer than greatest
width which occurs about midpoint of wing; apical one-fifth of wing mostly with solid
dark brown bands, most of hypostigmatic cell completely dark brown suffused; male
pilula axillaris with head about 3.OX wider than base of pedicel; male abdomen with
most tergites and sternites (especially toward posterior end) with numerous pore plates,
mainly associated with setal bases, giving abdomen a scaly appearance.
Material examined: ECUADOR. Guayas: Playas, II.26.1973, M. Deyrup (lm-FSCA),
Puerto Grande, Isla Puna, Gulf of Guayaquil, II.1936 Campos (-lm, 4f-FSCA); Posorja.
III Campos (2m lf-FSCA); Pt. Calera, IV.21,1968, E. Ball (If-FSCA).
PERU. Lambayaque. 40 mi. N. Chiclayo 1.17.1955, E. Schlinger & E. Ross (2m, CAS,
FSCA); Tumbes: Puerto Pizarro, 14 km. N. Tumbes, V.6.1959, W. Weyrauch (3f, IML;
Systematics: The broader wings and more extensive dark suffusion (Fig. 8) renders
this species distinctive in the genus. The male is distinguished from other species by
the abundance of tufted pore plates on the abdominal tergites giving the whole dorsum
of the abdomen a scaly appearance.


Contribution No. 698, Bureau of Entomology, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville, Florida.


BANKS, NATHAN. 1908. New tropical American Neuroptera. Proc. ent. Soc.
Washington 9: 30-34.
--. 1980. New genera and species of tropical Myrmeleonidae. J. New York Ent.
Soc. 17: 1-4.
--. 1909. New neuropteroid insects. Bull. Mus. Comp. Zool. (Harvard) 64:299-362,
7 Pls.
CAMPOS, R. FRANCISCO. 1922. Estudios sobre la fauna entomol6gica del Ecuador.
Revista colegio national Vicente Rocafuerte (Guayaquil) 4(7): 53-78.

September, 1989

Stange: New World Dimarini 461

ESBEN-PETERSEN, E. 1920. Revision of some of the type-specimens of Myr-
meleonidae described by Navas and placed in the Vienna Museum. Bull. Ann.
Soc. R. Ent. Belge 60: 190-196, Figs. 1-3.
HAGEN, HERMANN. 1860. Beitrag zur Kenntniss der Myrmeleon Arten. Stettin. Ent.
Ztg. 21: 359-369.
1861. Synopsis of the Neuroptera of North America, with a list of the south
American species, prepared in the Smithsonian Institution. Smiths. Misc. Coll.
4(1): i-xx + 1-347.
1866. Hemerobidarum Synopsis synonymica. Stettin. Ent. Ztg. 27: 369-462.
MARKL, WALTER. 1954. Vergleichend-morphologische Studien zur Systematik und
Klassifikation der Myrmeleoniden. Verh. Naturf. Ges. Basel 65: 178-263, Figs.
MCLACHLAN, ROBERT. 1867. New genera and species etc., of Neuropterous Insects;
and a revision of Mr. F. Walker's British Museum catalogue of Neuroptera. Part
II (1853) as far as the end of the genus Myrmeleon. J. Linn. Soc. (Zool.) 9 (1868):
230-281, P1. 8.
NAvAs, LONGINOS. 1912a. Myrm6leonides nouveaux ou peu connus. Ann. Soc. Sci-
ent. Bruxelles 36: 203-248 Figs. 1-3.
1912b. Notas sobre Mirmele6nidos. Broteria (Ser. Zool.) 10: 29-97.
1913. Nevropt6res. Mission du service geographique de l'armee pour la measure
d'un arc de meridien equatorial en Amerique du Sud sous le control scientifique
de l'Academie des Sciences 1899-1906. Paris: Gauthier-Villars 10(1): 68-77, P1. 4
(Figs. 1-10).
1914a. Neur6pteros nuevos o poco conocidos (Segunda series) Mem. R. Acad.
Cien. Artes Barcelona (3)11: 105-119, Figs. 1-6.
S1914b. Neur6pteros sudamericanos (Serie 1). Broteria (Ser. Zool.) 12: 45-56.
S1915. Neur6pteros nuevos o poco conocidos. (Serie 6). Mem. R. Acad. Cienc.
Artes Barcelona (3) 12(7): 119-136, Figs. 1-9.
1923. Alguns insects del Brasil. Rev. Mus. Paulista (Sao Paulo) 13: 767-774,
1 fig.
- 1933. Insectos suramericanos (6 Serie). Acad. Cienc. Nat. Madrid 29: 191-198,
Figs. 11-17.
1935. Insectos suramericanos (10 Serie). Rev. Acad. Cienc. Madrid 32: 360-
375, Figs. 56-62.
PERTY, MAXIMILLIAM. 1933. Delectus animalium articulatorum. Monachii: Arthor,
pp. i-iii, 1-222.
RIEK, E. F. 1967. Structures of unknown, possibly stridulatory, function on the wings
and body of Neuroptera; with an appendix on other endopterygote orders. J.
Australian Ent. Soc. 15: 337-348.
STANGE, LIONEL A. 1969. Myrmeleontidae of the Galapagos Islands. Acta Zool. Lil-
loana 25: 187-198, 15 figs.
.1984. Field Studies of the insect order Neuroptera in South America. Nat.
Geographic Soc. Research Reports (Washington, D.C.) 17: 51-76, 3 figs. 4 tables.
WALKER, FRANCIS. 1853. List of the specimens of neuropterous insects in the collec-
tion of the British Museum. Part II (Sialidae-Neompterides). London, pp. 193-
1859. Characters of undescribed Neuroptera in the collection of W. W. Saun-
ders. Trans. Ent. Soc. London 5(2): 176-199.

t 7

8 9








462 Florida Entomologist 72(3) September, 1989


Entomology-Biological Control
Florida A.&M. University
Tallahassee, FL 32307 U.S.A.

Department of Zoology
Panjab University
Chandigarh 160014 India


Bagous affinis Hustache is redescribed and its release in south Florida as a potential
biological control agent of Hydrilla verticillata is mentioned. A new species, B.
laevigatus O'Brien & Pajni, is described. Included are habitus photographs, specific
diagnoses, distributions, and line illustrations of the male phallus and endophallus and
of the female spermatheca and eighth sternum of both species.


Se redescribe Bagous affinis Hustache y se menciona su liberaci6n en el sur de la
Florida como agent potential para el control biol6gico de Hydrilla verticillata. Se
describe una especie nueva, B. laevigatus O'Brien & Pajni. Se incluyen fotografias del
aspect general, diagn6sticos especificos, distribuciones, y dibujos del falo y endofalo
de los machos y de la espermateca y del 8vo. estern6n de las hembras de ambas species.

Because Hydrilla verticillata (L. F.) L. C. has been designated the number 1 target
aquatic weed in the United States, research has been ongoing for several years to locate
biological control agents for this Old World plant. One such potential agent (Bagous
affinis Hustache) was discovered in Pakistan and India and a second species of Bagous,
new to science, was collected in the same areas. Both of these species attack tubers (or
turions) of hydrilla in soil exposed by receding waters. Some biology and field habits of
these two species were reported by Baloch, et al. (1980) as Bagous sp. nr. limosus
affiniss) and Bagous sp. (new species herein described). Buckingham (1988) has re-
ported the release of Bagous affinis in south Florida in April, 1987 by the U. S. Army
Corps of Engineers in a preliminary effort to control Hydrilla. He also discussed in
detail the history of the search for a control agent of this important aquatic weed.
No key exists to the 13 species of Bagous currently known from the Indian subcon-
tinent, but diagnoses of the two species herein and the illustrations included below will
distinguish the two Hydrilla tuber feeding species. A research study on the 30 Bagous
species (17 new to science) currently available from the Indian subcontinent should go
to press later this year and will include a key to all species. The new species is being
described herein to provide a name for publication of detailed life history studies that
have been conducted in quarantine at Gainesville, FL.

O'Brien & Pajni: Description of Hydrilla-feeding Weevils 463

Bagous affinis Hustache
(Figs. 1, 3, 5 & 6)

Bagous affinis Hustache 1926, p. 645.

Body medium sized, elongate-oval; sides weakly evenly rounded behind humeri to
declivity; black, with reddish brown antennal scape and funicle, tibial apex, and tarsi;
densely clothed with granulate, subcontiguous, pitted scales.
Male: Rostrum dorsally, evenly, strongly curved; ventrally evenly, weakly curved;
lacking carina; subcylindrical, ventral margin not angulate, nor carinate; basally trisul-
cate; basal 2/3 densely clothed with contiguous, pitted scales; apical 1/3 subglabrous;
apical 1/2 subquadrately expanded; 0.83 as long as prothorax. Head weakly convex;
densely clothed with subcontiguous, granulate, pitted scales; articulating area with
dense, whitish, plumose scales; frons moderately broad, 1.23 wider than apex of ros-
trum, strongly longitudinally sulcate; very weakly elevated along margin of eye, with
several, evident, suberect, curved setae. Antenna inserted at apical 1/3; scape long,
subclavate; funicle ca 1/10 shorter than scape; club broadly oval, subequal to funicular
segments 1-3 together. Prothorax 0.92 as long as broad; sides unevenly, weakly
rounded to strongly constricted apical 1/6; disc undulately convex with strong median
longitudinal sulcus, deeper and broader subapically and subbasally; basal margin lacking
transverse carina; with dense, coarse, rugulose coating of granulate, imbricate, pitted,
sublunate, distinctly emarginate scales; color dark blackish brown, with 3 uneven, inter-
rupted longitudinal paler whitish brown vittae. Elytra with well developed obliquely
rounded humeri; markedly wider than prothorax, odd intervals convex, with rows of
conspicuous, subrecumbent, pale setae; even intervals flat; lacking swelling or tubercle
in front of declivity; 5th interval with weakly developed, though evident, subangulate
declivital callous; striae distinctly grooved with moderately elongate, evident punctures,
densely clothed with contiguous, pitted scales; color dark brown to black, with numer-
ous, scattered, small, paler brown to whitish maculae; very strongly narrowed behind
declivity to non-acuminate, scarcely emarginate apices. Prosternum deeply broadly
sulcate; sides of sulcus subparallel, scarcely narrower at apex, sharply raised, markedly
rounded, just in front of coxae. Abdominal sternum 1 broadly deeply impressed; ster-
num 2 flat; sternum 5 laterally strongly impressed, medium area weakly impressed,
with pair of evident, small, subapical tubercles; sternum 1 ca 1/3 longer than 2, 5
subequal to 2. Legs moderately long; femora strongly clavate; tibiae distinctly bisinuate,
moderately stout, inner surface not denticulate, with several moderately short, evident
bristles, apex not distinctly narrowed, strongly curved inward very near apex; tibial
uncus stout, shorter than width of tibiae; tarsi moderately short, not linear, segments
1-3 widened towards apex; segment 4 subequal to 2 and 3 together; ventrally with
numerous, apical, fine bristles and apical pad of fine pubescence. Length, pronotum and
elytron: 2.3 mm.
Female: Very similar to male except: Rostrum lacking evident median sulcus, me-
dially flattened. Abdominal sternum 1 basally, weakly, broadly impressed, apical half
weakly convex; sternum 5 flattened medially, lacking evident subapical tubercles.
Length, pronotum and elytron: 3.6 mm.

Remarks and Comparative Notes.-The 284 specimens on hand range in size from 2.8
to 4.0 mm. In India and Pakistan, this relatively nondescript species superficially resem-
bles the smaller (usually less than 2.5 mm.) vicinus Hustache, but the latter has round,
granulate, not emarginate scales and its scale color is mainly brown with whitish
maculae and vittae and some black areas, not mainly black with whitish and brownish
maculae and whitish prothoracic vittae as in affinis. Most affinis are black and grey in
color and have scarcely projecting, round to angulate, declivital calli.

Florida Entomologist 72(3)




Fig. 1. B. affinis, male; Fig. 2. B. laevigatus, male; Fig. 3. B. affinis, female; Fig.
4. B. laevigatus, female, a, phallus, dorsal view; b, endophallus, [Fig. 1] dorsal view
only, [Fig. 2] left, dorsal view; right, lateral view; c, phallus, lateral view; d, sper-
matheca, lateral view; e, 8th sternum, ventral view. Lines = 1mm. (line A, males; line
B, females).

In the United States, this species most resembles texanus Tanner, and keys to this
species in couplet 7 in his North American revision (Tanner, 1943) but texanus has
round, not emarginate, thoracic scales and has shallow, small, poorly defined, subapical,
and subbasal, median thoracic impressions.
Type locality.-India, Uttar Pradesh State, Sarda River.
Range.-Known from Bangladesh, India, Pakistan, Thailand, and introduced in south
Florida, U.S.A.
Material Examined.-BANGLADESH: Rajshahi: Dinajpur, X-1969 (3), X-1970 (2)
Barbe. INDIA: Bihar: Pusa, at light, 5-VII-1935 (1) H. S. Pruthi. Karnataka: Banga-
lore, at UV, V-1982 (9) J. K. Balciunas, M. C. Minno, Light Traps & Aquatic Plants,
MCM-82-1, V-VI-1982 (17) M. C. Minno, J. K. Balciunas, Hydrilla Tests, V-VI-1982 (8)
J. K. Balciunas, C. Minno; 40 km. W. Bangalore, Bombay Rd., [no ecological data] (10),
ex shore washed debris (2), at night hand picked from flowers & leaves of Potamogeton
(4), 4-X-1985, [no ecological data], 7-X-1985 (4) C. W. & L. B. O'Brien; 35.5 km. W. S.
W. Bangalore, Dasappadoddi Pond, at UV, No. KAR82BL3, 24-V-1982 (14) J. K. Bal-
ciunas & M. C. Minno; Bangalore, Hoskote Rd., Avaleahalli, at UV, CAB 85-19, 15-V-


September, 1989



O'Brien & Pajni: Description of Hydrilla-feeding Weevils 465

1985 (64) C. A. Bennett & G. R. Buckingham; 18 km. E. Bangalore, Madras Rd., [no
ecological data] (2), at [white] light (5), under stones (1), UV (12), 2-X-1985 C. W. &
L. B. O'Brien; Bangalore, Mysore Rd., Blacklight, FBCL-83-1005 (6), FBCL-83-1007
(7), V-1983, light traps, FBCL-83-1018 (13), FBCL-83-1019 (4), VII-X-1983 (3) coll.
CIBC; [parent stock], Bangalore, VIII-X-1983, [offspring], FL., Alachua Co., Gaines-
ville, Lab Colony, reared on tubers Hydrilla verticillata (L. F.) Royle (1) G. R. Buck-
ingham; Magadi, Adults on Hydrilla, shore and submerged, FBCL-85-1003, 1 & 7-I-
1985 (1) Coll. CIBC-IS, FBCL 83-1015 (38), FBCL 83-1014 (24), VII-VIII-1983 [CIBC-
IS]; near Magadi, XI-1984 (4) K. N. Nair. Maharashtra: Pashan Lake, Pune, 18-X-1985
(1) C. W. & L. B. O'Brien. PAKISTAN: Punjab: Akhori, adult hibernating under dry
plant of Hydrilla verticillata, 11-II-1972 (7) [no collector]; Nandipur, larvae boring
tuber of Hydrilla verticillata, 6-IV-1974 (5), 5-V-74 (2) [no collector]; 10 km. N. Gujran-
wata, coll. in soil nr. Hydrilla verticillata (L. F.) Royle tubers, CAB-85-5.1, 23-IV-1985
(2) [no collector]. THAILAND: Kaen Municipality, 27-V-1954 (1) R. E. Elbel.

Bagous laevigatus O'Brien and Pajni, new species.
(Figs. 2, 4, 7 & 8)

Body medium sized, elongate-oval; sides subparallel; black, and reddish black, with
reddish brown antennal funicle, tibiae and tarsi; densely clothed with contiguous and
imbricate, flat, finely pitted or not pitted, black, and grey scales.
Holotype male: Rostrum moderately unevenly curved, moderately depressed in api-
cal 1/2; lacking carina; with long, evident, lateral sulcus; subcylindrical, ventral margin
not angulate, nor carinate; densely clothed with contiguous, subgranulate scales; apical
1/3 strongly quadrately expanded; 0.80 as long as prothorax. Head moderately convex,
densely clothed with subgranulate, contiguous scales; frons moderately broad, 1.31
wider than apex of rostrum, medially flattened with large, deep, median fovea, slightly
elevated along margin of eye, elevation with several long, erect, curved setae. Antenna
inserted at apical 1/3; scape long, subclavate; funicle ca 1/5 shorter than scape; club
broadly oval, subequal in length to funicular segments 1-3 together. Prothorax 0.94 as
long as broad; sides subparallel, widest near base, weakly narrowed to rounded apical
1/5, there weakly constricted; disc moderately convex, with indistinct, very narrow,
median, longitudinal impression, and small, broader, shallow, basal impression; basal
margin lacking transverse carina; densely clothed with contiguous, smooth, round to
oval scales and several sparse, scarcely distinct punctures, and numerous, fine, scarcely
evident, apical and marginal, suberect setae; black, with median, narrow, basal, grey
fascia and pair of lateral, broader, basal, grey maculae, and several indistinct, small,
scattered, grey maculae. Elytra with obliquely, weakly angled, rounded, moderately
developed humeri; markedly wider than prothorax; intervals flat; lacking swelling or
tubercle in front of declivity; striae distinctly, finely grooved, with scarcely evident
small elongate punctures; 5th interval with weak, scarcely evident, declivital callous;
color black with numerous, scattered, small, grey maculae, and with one medium sized
macula anterior to declivity on intervals 3 and 4; sides subparallel, weakly roundly
expanded behind humeri to declivity, there suddenly narrowed to non-acuminate,
weakly emarginate apices. Prosternum broadly, moderately shallowly sulcate; sides of
sulcus moderately biangulate, narrowest at apical constriction, lacking sharp margins,
moderately rounded in front of coxae. Abdomial sternum 1 broadly, deeply impressed;
2 flattened, declivous at apical 1/10; sternum 5 scarcely impressed laterally, remainder
flattened; sternum 1 ca 1/5 longer than 2, 2 ca 2/5 longer than 5; sterna 3 and 4 slightly
shorter than 5. Legs moderately short; femora strongly clavate; tibiae stout, sinuate,
moderately curved inward apically, inner margin with several small evident denticles,
and moderately long bristles, apex not distinctly narrowed; with long tibial uncus ca as

Florida Entomologist 72(3)

Figs. 5 & 6. B. affinis, Figs. 7 & 8. B. laevigatus.


September, 1989

O'Brien & Pajni: Description of Hydrilla-feeding Weevils 467

long as width of tibia; tarsi moderately long, sublinear, 3rd segment not wider than
2nd, 4th segment as long as 2 and 3 together, ventrally with several long, apico-lateral
bristles, no pubescent pads. Length, pronotum and elytron: 3.8 mm.
Allotype female: Very similar to male except: Prothorax 0.87 as long as broad.
Abdominal sternum 1 broadly, very shallowly impressed; sternum 5 with strong lateral
impression. Length, pronotum and elytron: 4.2 mm.

Etymological Note.-This epithet is from the Latin, laevigatus (= smooth, slippery),
and refers to the extremely smooth and shining surface of this species.

Remarks and Comparative Notes.-The 783 specimens on hand holotypee, allotype
and 781 paratypes) range in size from 2.6 to 4.6 mm. This distinctive, smooth, and shiny
species is not likely to be confused with any other from the Indian region or Florida.
Superficially it resembles affinis Hustache, but lacks its deep prothoracic impressions,
and coarsely pitted, emarginate scales, and laevigatus has scarcely evident declivital
calli. Both brown, and grey color forms are common.

Type locality.-India, Karnataka State, 18 km. E. Bangalore, Madras Road.

Range.-Known from India and Pakistan.

Material Examined.-INDIA. Delhi: Azadpur Colony, at Light, 16-V-1959 (31) Balder
Pd. Coll.; Kallaspuri Palem, at Light, 4-VII-1981 (1) Maya Ram Coll. Karnataka: Ban-
galore, at UV, MCM-82-1 (5), Light Traps & Aquatic Plants (3), V-VI-1982 M. C.
Minno, J. K. Balciunas; [parent stock] 40 km. W. Bangalore, Bombay Rd., ex tuber
Potamogeton pectinatus [in lab], V-VII-1983 (9) [CIBC-IS], [offspring] FL., Alachua
Co., Gainesville, F4, reared on tubers Hydrilla verticillata (L .F.) Royle, USDA Lab
Colony (1) G. R. Buckingham, [no ecological data] (13), ex shore washed debris (21),
4-X-1985 C. W. & L. B. O'Brien; 35.5 km. W. S. W. Bangalore, Dasappadoddi Pond,
at UV, KAR82BL1, 17-V-1982 (1), 24-V-1982 (4) J. K. Balciunas & M. C. Minno; Ban-
galore, Hoskote Rd., Avaleahalli, at UV, CAB-85-19, 15-V-1985 (82) C. A. Bennett &
G. R. Buckingham; nr. Hoskote, nr. Bangalore, Hydrilla verticillata on shore and
submersed, FBCL-84-1014-1016, 16-VII-1984 (4) CIBC-IS; 18 km. E. Bangalore, Mad-
ras Rd., [no ecological data] (10), UV Light holotypee + allotype + 6), UV (461), at
[white] light (33), under stones (22), under live Polygonum spp. & dead Hydrilla in dry
lake (27), 2-X-1985 C. W. & L. B. O'Brien, at [white] light (227), to white sheet at dusk
(4) L. B. O'Brien; Bangalore, Mysore Rd., light trap, FBCL-83-1018 (2), FBCL-83-1019
(2), 1983, FBCL-83-1007, V-1983 (2) [CIBC-IS]; Magadi, Adults on Hydrilla, shore and
submerged, FBCL-85-1003, 1 & 7-1-1985 (3) Coll. CIBCIS, collected on Hydrilla ver-
ticillata (L. F.) Royle, FBCL-83-1014 (23), FBCL-83-1015 (27), VII-VIII-1983 Coll.
CIBC. Punjab: Chandigarh, tube light, 10-VII-1977 (2) [no collector]; Chandigarh, Rail-
way Station, Tube Light, 10-VII-1977 (3), 25-VII-1977 (1) [no collector]; Kaithal, Light,
7-V-1977 (1) [no collector]. PAKISTAN: Punjab: Akhori, CIBC Hyd. 2/72-7, 11-II-1972
(2) [CIBC-IS]; Lahore, VII-VIII-1957 (1) J. Maldonado. Sind: nr. Karachi, Debris on
beach, "INDIA, 1952", BM 1952-143, 16-XII-1951 (1) T. Clay; Haleji Lake (Indus Delta),
in shore washed debris, mostly Hydrilla, 23-VIII-1985 (1) C. W. & L. B. O'Brien;
Keenjhar Lake (Indus Delta), in shore washed debris, mostly Hydrilla, 24-VIII-1985
(1) C. W. & L. B. O'Brien; Thatta, at light, 15-X-1971 (1) S. Khan; Sukkur, Adult
hibernating under dry plant of Hydrilla verticillata, CIBC Hyd. 2/72-14, 1423, 13-II-
1972 (3) [CIBC-IS].
The holotype, allotype and numerous paratypes are deposited in the senior author's
collection. Paratypes are deposited also in the following collections: Auburn University,
Auburn, AL; Bombay Natural History Society Research Centre, Bharatpur, India;

Florida Entomologist 72(3)

British Museum (Natural History), London, England; California Academy of Sciences,
San Francisco, CA; California State University, Long Beach, CA; College of Madura,
Madurai, India; Commonwealth Institute of Biological Control, Bangalore, India; Com-
monwealth Institute of Biological Control, Islamabad, Pakistan; Florida State Collection
of Arthropods, Gainesville, FL; Indian Agriculture Research Institute, New Delhi,
India; Kerala Agricultural University, Trichur, India; Modern College, Pune, India;
Museum National d'Histoire Naturelle, Paris, France; Panjab University, Chandigarh,
India; Staatliches Museum fur Tierkunde, Dresden, GDR; Texas A&M University, Col-
lege Station, TX; United States National Museum, Washington, DC; and Zoological
Survey of India, Calcutta, India.


We wish to express our special thanks to Dr. Gary Buckingham, USDA-ARS,
Gainesville, FL, who made available to us the majority of the specimens used in this
study and who aided the senior author in obtaining funding for the travel needed to
complete the research.
We also thank the following individuals and their institutions for the loan of speci-
mens and/or assistance during the senior author's field work in India and Pakistan: Prof.
Imtiaz Ahmad, Department of Zoology, University of Karachi, Karachi, Pakistan; Dr.
John George M., Bombay Natural History Society Research Centre, Bharatpur, India;
Drs. Swaraj Ghai and V. V. Ramamurthy, Division of Entomology, Indian Agricultural
Research Institute, New Delhi, India; Dr. Peediakal J. Joy, Regional Agricultural Re-
search Station, Kottayam, India; Dr. R. Krause, Staatliches Museum fur Tierkunde,
Dresden, GDR; Dr. A. Ikram Mohyuddin, Commonwealth Institute of Biological Con-
trol, Rawalpindi, Pakistan; Dr. S. Y. Paranjape, Department of Zoology, Modern Col-
lege, Pune, India. Mr. Alexander Riedel, Friedberg, FRG; Drs. T. Sankaran and M.
J. Chacko, Commonwealth Institute of Biological Control, Bangalore, India; Drs. K. G.
Sivaramakrishnan and K. Venkataraman, Department of Zoology, College of Madura,
Madurai, India; Mr. Richard Thompson, Department of Entomology, British Museum
(Natural History), London, England; and Dr. B. K. Tikader, Zoological Survey of India,
Calcutta, India.
This research was supported in part by a travel grant from the U. S. Army Corps
of Engineers to study types in London and Paris; and a travel grant to Pakistan and
India from the U. S. Department of Agriculture, Office of International Cooperative
Development, to collect Bagous and to obtain host data. Research was also partially
supported by a research program (FLAX 85006) of the SEA/CR, USDA.


BALOCH, G. M., SANA-ULLAH, AND M. A. GHANI. 1980. Some promising insects for
the biological control of Hydrilla verticillata in Pakistan. Trop. Pest Manage.
26(2): 194-200, illus.
BUCKINGHAM, G. R. 1988. Reunion in Florida Hydrilla, a weevil, and a fly. Aqua-
tics 10(1): 19-25, illus.
HUSTACHE, A. 1926. Nouveaux Bagous d'Asie (Coleoptera, Curculionidae). Ann.
Mag. Nat. Hist. 17: 641-647.
TANNER, V. M. 1943. A study of the subtribe Hydronomi with a description of new
species, (Curculionidae) study No. VI. Great Basin Nat. 4(1-2): 1-38, illus.


September, 1989


v -m


Howard & Solis: Mahogany Webworm 469


University of Florida
Institute of Food and Agriculture
Fort Lauderdale Research and Education Center
3205 College Avenue
Fort Lauderdale, Florida 33314

Department of Entomology
University of Maryland
College Park, Maryland 20742


The life history and host plant relationships of the mahogany webworm, Macalla
thyrsisalis Walker (Lepidoptera: Pyralidae: Epipaschiinae), were investigated in
Florida and the distribution of this species in the Americas was determined by examin-
ing museum specimens in the U.S. National Museum of Natural History.
Macalla thyrsisalis is distributed throughout the natural range of American
mahoganies, Swietenia spp., and some localities where mahoganies have been intro-
duced. In Florida, natural infestations have been observed only on native West Indies
mahogany, S. mahagoni Jacquin, but larvae feed on leaves of some close relatives of
this species if placed on them. Honduras mahogany (S. macrophylla King) and Spanish-
cedar (Cedrela odorata L.) are suspected to be hosts in Central and South America.
In Florida, the adult moths fly in early spring. Larvae spin webs and feed on
mahogany leaves during the ca. 5-week period of the spring flush, then leave the trees
to pupate. Larfal populations greatly diminish by early summer due to unknown factors.
Parasitoids include Lespesia n. sp. (Diptera: Tachinidae), reared from 8.8% of 25
mahogany webworms, and two species of Braconidae: Habrobracon sp. near hebetor
Say and Apanteles sp. Apparent relationships between mahogany webworm popula-
tions, spring leaf flush of mahoganies and photoperiodic and climatic factors are dis-


En la Florida, E. U. A., se investig6 la historic natural de Macalla thyrsisalis
Walker (Lepidoptera: Pyralidae: Epipaschiinae) y sus relaciones con plants hospederas,
y se determine la distribuci6n de esta especie en las Americas examinando ejemplares
en el Museo Nacional de Historia Natural de Los Estados Unidos.
Macalla thyrsisalis esta distribuido por toda la habitaci6n natural de las caobas
americanas, Swietenia spp., y en algunas localidades donde se han introducido las
caobas. En la Florida, se han observado infestaciones naturales de M. thyrsisalis sol-
amente en la caoba antillana, S. mahagoni Jacquin, la cual es native de esta area, sin
embargo las larvas se alimentan de las hojas de species pr6ximas cuando se colocan en
ellas. Se supone que la caoba Hondurefa (S. macrophylla King) y el cedro Espahol
(Cedrela odorata L.) son sus plants hospederas en Centroambrica y Sudambrica.
En la Florida, los adults vuelan al principio de la primavera. Las larvas hacen su
tejido telearafoso y se alimentan de las hojas de la caoba durante el period de la
brotadura de primavera (alrededor de cinco semanas) y despu6s dejan los arboles para
pupar. Las poblaciones larvales disminuyen antes de los principios del verano debido a
factors todavia desconocidos.

Florida Entomologist 72(3)

Los parasitoides incluyen Lespesia n. sp. (Diptera: Tachinidae), criado de 8.8% de
125 larvas, y dos species de Braconidae: Habrobracon sp. cerca de hebetor Say y
Apanteles sp.
Se discuten las relaciones aparentes entire las poblaciones de M. thyrsisalis, la
brotadura de primavera de las caobas y los factors climaticos y de fotoperiodo.

West Indies mahogany. Swietenia mahaxoni Jacauin. (Meliaceae) is native to south-

Fig. 1.
Fig. 1 and 2. (1) Mahogany webworm, Macalla thyrsisalis adult moth (2) Mahogany
webworm larva.

September, 1989


Howard & Solis: Mahogany Webworm

Fig. 2.

ern Florida, the Bahamas, and the Greater Antilles except Puerto Rico (Pennington et
al. 1981). In southern Florida, where it is a preferred tree for landscaping (Morton
1987), there is public concern each spring over the appearance of a caterpillar, the
mahogany webworm, Macalla thyrsisalis Walker (Lepidoptera: Pyralidae: Epipas-
chiinae), which produces extensive webbing and consumes foliage of these trees
(Chellman 1978, Reinert & Howard 1982) (Fig. 1 & 2).
The present paper is the first report on the life history and host plant relationships
of mahogany webworm and its distribution in the Americas.


Specimens of Macalla in the U.S. National Museum of Natural History, Smithsonian
Institution (USNM), were examined and genitalia and heads dissected to identify M.
thyrsisalis. The distribution of the species was determined from collection data.
To identify the host plants other than West Indies mahogany in Florida, trees in the
vicinity of infested West Indies mahoganies were often examined during the webworm
season. Several times, mahogany webworms were transferred from West Indies
mahoganies, which they had naturally infested, to other members of the Meliaceae,
including Honduras mahogany (S. macrophylla King), Spanish-Cedar (Cedrela odorata
L.), African mahogany (Khaya sp.), pimientilla (Trichilia trifolia L.) and China-berry
(Melia azedarach L.), and observed for several days to determine whether they would
feed on them.
To determine when adults were active in Florida, daily collections were made from
January 1985 through December 1987 from two blacklight (BL) traps ca. 200 m apart

Florida Entomologist 72(3)

each near a West Indies mahogany at the Fort Lauderdale Research & Education
Center (FLREC). These standard design traps, i.e., similar to those of Mahrt et. al.
(1987), used 15 watt BL bulbs. Insects entering a trap fell through a funnel into a water
Twigs and foliage of mahoganies were examined for eggs when the adult mahogany
webworm moths were flying (based on BL collections) and prior to the appearance of
webworms on foliage. Also, leaves and branches with early instar webworms were
scrutinized. About 15 hours were devoted to this search. Moths were reared and kept
in cages of various sizes with mahogany foliage and various nectar sources or sucrose
solutions and the foliage examined frequently for eggs.
Casual observations had indicated that the presence of mahogany webworms on
foliage coincided with the flush of new leaves in the spring (Reinert & Howard 1982).
To define this period, observations were made on 7 trees in a planting of 30 West Indies
mahoganies on the grounds of the FLREC. These trees were 3 years old at the begin-
ning of the study. Heights of the trees were measured each June from 1983 1987.
During March 1985 to November 1987, these trees were observed at least weekly
for webworms, and beginning in April each year they were observed every 1-3 days
until the first larval webworms of the season were observed. When present, a count of
the number of webworms / tree was made. At each observation, each tree was classified
based on the stage of development of most of its foliage into one of the following
categories: (1) mostly very young leaves petioless and leaf blades deep reddish in color),
(2) mostly expanding leaves (leaf blades light green, tender), or (3) mostly fully ex-
panded, hardened-off leaves (deep green, relatively stiff and tough). Temperature and
precipitation data were obtained from a weather station operated at the FLREC.
West Indies mahoganies also were examined for webworms, webs, or feeding dam-
age during occasional field trips to various sites in southern Florida during 1983-1987.
To determine the duration of the larval stage, 15 field- collected mahogany web-
worms 2-4 mm long, the smallest size that we have observed and presumedly the first
instar, were placed three / plant on each of five West Indies mahogany seedlings ca. 50
cm tall in an outdoor cage in April 1985. Maximum and minimun temperatures and the
lengths of the larvae at rest were recorded daily. The number of webworms on plants
at each observation diminished gradually, possible due to dispersal or predators, so that
after 11 days, only four webworms were still on the plants. These were placed individu-
ally in Petri dishes with mahogany foliage and kept in the same outdoor location until
adult eclosion.
To identify natural enemies of mahogany webworm, in May- June, 1986, 120 late
instar larvae were collected from various sites in southern Florida from Key Largo to
Fort Lauderdale, reared on mahogany foliage until pupation or the emergence of
parasitoids, then kept in rearing cups until emergence of adult moths or parasitoids.


Distribution. Collecting data of specimens of adults and larvae identified as M. thyr-
sisalis in the USNM were as follows: FLORIDA: Broward Co.: 24-III-1986, 12-IV-1986,
4, 7, 14, 18, 20, 28-V-1986, 14-VI-1986, 30-IV-1987, F. W. Howard; Ft. Lauderdale,
1985, Swietenia mahagoni, F. W. Howard; Dade Co.: IV-1954; 23-V-1945 ex. mahogany;
Coconut Grove, (Swietenia mahagoni), Tuthill; Coral Gables, 4-V-1946, Mahogany, G.
B. Merrill; Homestead: 16-V-1982, C. V. Covell, Jr.; 19-IV-1944, mahogany on leaves,
G. B. Merrill; Miami: 29-V-44, ex mahogany; 21-IV-1944, defoliating mahogany trees,
G. B. Merrill; 10 April 1945, mahogany, G. B. Merrill; 24 April 1944, mahogany tree,
Tuthill; 29-V-1944, ex mahogany S. S.; 5-V-1966, Swietenia mahagoni (W. I.
mahogany), D. H. Habeck; BAHAMAS: Freeport, 20-27-VI-1987, W. E. Steiner, M. J.

September, 1989

Howard & Solis: Mahogany Webworm 473

S- - -Tropc of Cancer

Swletenia mahagoni

.......TT___%ogc _? rorn. __-n___

Fig. 3. Localities where mahogany webworms, Macalla thyrsisalis have been col-
lected (distribution of West Indies and Honduran mahogany after Pennington at al.

and R. Molineaux; TRINIDAD: E. W. Rorer Coll; Caparo, XII-1905 (S. M. Klages);
Curepe, 29-X-1979, M. J. W. Cock; DOMINICAN REPUBLIC: El Seibo Province, 15 km.
s. Miches, ca. 500 m, 31 V-1973, Don and Mignon Davis; La Vega Province, Constanza,
1164 m, Hotel Nueva Suiza, 29-V-1973, Don and Mignon Davis; MEXICO: Cordoba, V-
'06, '08, Fred K. Knabb, Collector; GUATEMALA: Tikal, 200 m, 10-V-1960, S.
Steinhauser; HONDURAS (Type series at British Museum, Natural History); COSTA
RICA: La Selva Biological Station, Puerto Viejo de Sarapiqui, Pr. Heredia, 40 m, III-
1986, M. M. Chavarria; Sirena, Corcovado National Park, Osa Peninsula, 5- 11-1-1981,
D.H. Janzen and W. Hallachs; PANAMA: Portobelo, 11-III, August Busck; BRAZIL;
Para, Belem, 20 m, i. 1984, V.O. Becker.
Thus, the distribution of M. thyrsisalis coincides in general with the natural distri-
bution of West Indies mahogany in Florida, the Bahamas and the Greater Antilles and
that of Honduras mahogany in Mexico, and Central and South America as reported by
Pennington et al. 1981 (Fig. 3). Islands of the Bahamas other than Grand Bahama,
islands of the Greater Antilles other than Hispaniola, and inland areas of South America
were major areas within the natural range of mahoganies not represented by specimens
of M. thyrsisalis. Belem, Brazil, and Trinidad fall outside of the natural range of Hon-
duras mahogany, but within the natural range of Spanish-cedar, and Honduras
mahoganies have been introduced into these localities (Lamb 1966, P. Celestino Filho,
Empresa Brasileira de Pesquisa Agropecuaria, Belem, Brazil, personal communication).
Most of the collection dates were in April and May.
Specimens of the closely related M. hyalinalis complex have been collected in Puerto

474 Florida Entomologist 72(3) September, 1989

Rico, Cuba, Jamaica, Mexico, Honduras, El Salvador, Costa Rica, Panama, Venezuela,
Guyana, Brazil and Argentina. This group has been observed feeding on Cedrela sp. in
Costa Rica (D. Janzen, University of Pennsylvania, personal communication) and on
Spanish-cedar in Tamaulipas, Mexico (M. A. Solis).
Hosts. We have observed natural infestations of mahogany weborm in Florida on
West Indies mahoganies, but not other plant species, including 20 Honduras
mahoganies, 6 Spanish-cedars and 4 African mahoganies in the vicinity of infested West
Indies mahoganies. Mahogany webworms transferred from West Indies mahoganies to
these species made webs and fed on the leaves. However, they did not feed when
transferred to leaves of China-berry or pimientilla.
Life history. One to a few mahogany webworm moths were caught in the BL traps
at the FLREC on each of the following days: March 14, 21, 27, 30 (1985); April 11, 12,
14, 21, 29, May 28, 30, June 2, 4 (1986); April 23, 24, 27, 29, 30, May 4, 5, 6, 7, 8, 14,
18, 20, 28, June 22 and July 7 (1987). Based on these data, flight of the moths occurred
each spring, during a short early period in 1985, a later and more prolonged period in
1986, and a somewhat later period in 1987 (Figs. 4 6). If moths flew at other times of
year, they were not detected by our methods.
We did not observe eggs of mahogany webworms. Peterson (1963) described eggs
of this species obtained by confining female moths in plastic bags as "ovoid, exceedingly
pliable, near white in color and very adhesive. Within a given mass the eggs under
pressure assume varying shapes". This reported pliability and an unusually long
ovipositor in the adult female suggests that the eggs may be inserted, e.g., into bark
crevices, and therefore difficult to observe in the field.
Larvae are solitary. Each spins a single web which pulls several leaves together and
feeds partially concealed within it, although in some cases, larvae construct webs in
close proximity so that it may appear that two or more larvae occupy a single web.
A few webworms were seen on one of the seven trees under observation in spring
of 1984. By spring of 1985, all seven trees were infested. The mean height in meters of
the trees in successive years from 1983 to 1987 as as follows: 1.49, 1.63, 2.46, 3.11 and
4.12. After 1985 there was no increase in the webworm populations corresponding to
the increase in tree size.
The larval populations were present during the period of the spring flush and leaf
expansion, with a short early period in 1985, a somewhat more prolonged period in 1986,
and a shorter and later period in 1987. Thus, both the period of adult flight and presence
of larval populations appeared to have an imprecise coincidence with the spring flush
(Figs. 4 6). The first larvae each year were always on very young leaves, and larvae
that were transferred to new trees or young seedlings invariably moved to the newest
leaves and built webs and fed. Populations fell off as the spring leaves matured and
hardened. The trees continued to grow and produce new leaves throughout the summer,
however, and thus appeared to remain susceptible to webworm attack. Except for
single larvae seen on July 21, 1986 and July 16, 1987, mahogany webworms were never
seen in the field after June of each year. The webworm collected as an apparent first
instar larva on July 21, 1986 and kept in the laboratory pupated on August 4 and closed
as an adult on September 5. Thus, mahogany webworms may be present in very low
population densities on mahoganies after June of each year.
The observations on the trees at the FLREC were consistent with observations on
occasional field trips throughout southern Florida during 1983-1987, i. e. West Indies
mahoganies are often infested with webworms during spring flush and leaf expansion
in April-June, but not at other times of year.
Of the 5 apparently first instar webworms placed on mahogany seedlings on April
19, 1985 to determine the duration of life stages, four remained on the plants after 11
days. A daily attrition of the population was observed during these 11 days, possibly

Howard & Solis: Mahogany Webworm






Fig. 4.

I ;;_j

Florida Entomologist 72(3)


Fig. 5.

Figs. 4 6. Population patterns in Macalla thyrisalis and the spring flush in seven
West Indies mahogany trees at Ft. Lauderdale, Florida during spring months of 1985,
1986 and 1987. Portion of trees with (a) older leaves, (b) mostly newly opened leaves,
(c) young, partly expanded leaves, (d) new, fully expanded leaves; (e) period during
which adult moths were captured in blacklight traps, and (f) total numbers of larvae on
seven trees.

due to predators or dispersal from the seedlings. Webworms attained a maximum length
at rest of ca. 24mm and then entered a prepupal stage for 5 7 days, during which they
built cocoons. The pupal period lasted 18 20 days. Three adult moths closed on May
28 and 29. During these observations, the average minimum and maximum tempera-
tures were 19.60 and 34.90, respectively, this being typical of the warmer months in
southern Florida.

September, 1989

Howard & Solis: Mahogany Webworm



Fig. 6.
In late spring it is common to see mature mahogany webworms crawling on the
ground beneath mahogany trees, presumedly searching for pupation sites. We found no
mahogany webworm cocoons in lawns in the vicinity of mahoganies and only one cocoon
in a bark crevice of a mahogany tree. Mahogany trees grown in our nursery are infested
each year by webworms, and in late spring and summer it is common to find the cocoons
in spaces between the soil and containers or beneath the containers. Larvae reared on
seedlings in a cage pupated beneath a layer of sand on the bottom.
Natural enemies. Of 125 pupae collected from various sites in southern Florida
between May 7 and June 11, 1986, and kept in separate rearing cups, 47.2% closed as
adult moths during June and July. A fly, Lespesia n. sp.(Tachinidae) (ident. N. E.
Woodley, U.S.D.A. and B. E. Cooper, Agriculture Canada) emerged from 8.8% of the
pupal cases. These pupae had been reared from larvae collected in Fort Lauderdale,
Coral Gables, Opa-locka, and Key Largo. Habrobracon sp. near hebetor Say and Apan-

478 Florida Entomologist 72(3) September, 1989

teles sp. (Braconidae) (ident. P. M. Marsh, U.S.D.A.) were reared from mahogany
webworms collected from Fort Lauderdale and Key Largo. The larvae of these
braconids emerged from the later instar webworms and formed cocoons in the webbing
from which the adults were reared. Similar cocoons were often seen in webbing pro-
duced by mahogany webworms in the field. Since adults failed to eclose from 44.0% of
the 125 mahogany webworm pupae by August 11, these were dissected. Only 4.8% were


Based on this study, the life history of the mahogany webworm in Florida is as
follows: the adult moths fly in early spring. Oviposition sites are not yet known. The
larval stage lasts just over 10 days or less. Webworm populations build up during a
period of about 5 weeks that coincides with the spring flush and expansion of new
leaves. The mature larvae pupate in bark crevices or protective sites on the ground.
Braconid wasps and tachinid flies parasitize immature stages. Under laboratory condi-
tions, almost half of the pupae closed in June or early July. If this were true in the
field, we would expect that adults would continue to be caught in BL traps and larvae
would continue to infest foliage at least throughout the summer, but this was not the
case. Perhaps the adults are inactive until the following spring, or migrate or switch to
host plants other than West Indies mahoganies.
Although the insect is widespread in southern Florida, severe mahogany webworm
infestations are very unevenly distributed, and tend to be concentrated in the same
areas each year. Dense populations are most often observed on mature mahogany trees
in extensive uniform street plantings. Populations of mahogany webworms are not
uniformly distributed on individual trees of infested plantings. This appears to be at
least partly attributable to variability in timing of the spring flush in individual trees.
Although mahogany webworms attacked the 10 young trees under our close observa-
tion, and populations did not increase correspondingly as the trees grew larger, in
general, solitary and younger mahogany trees tend to be free of this insect. These
observations suggest that individual West Indies mahogany trees vary in susceptibility
to the webworms and that the insects are relatively inefficient in finding new host trees.
Maintaining West Indies mahoganies among a diversity of other tree species probably
tends to prevent mahogany webworm attack.
Peaks of flushing in tropical trees often occur at or near equinoxes (Alvim 1964), as
is true of West Indies mahogany, suggesting that flushing is influenced by the photo-
period. Broschat & Donselman (1983) showed that growth of West Indies mahoganies
is influenced by temperature as well as photoperiod. There was a delay in the spring
flush and in spring flight of adults in 1987. This may have been associated with cool
temperatures during or previous to that period, as the mean minimum temperature in
April 1987 was 14.3 C. compared to 17.2 C. in April of 1985 when leaf flush and the
webworm population occurred earlier in the spring.
A common assumption is that flushing in trees takes place in response to rainfall,
but in many tropical tree species this is not the case and in some species flushing even
occurs during the dry season, sometimes just prior to the rainy season (Longman &
Jenik 1974). We detected no association of flushing with rainfall in the 10 trees we
observed for foliage development. In fact, the 4-month period prior to the relatively
early spring flush in 1985 was comparatively dry with a total precipitation from January
through March of 76.9 mm. In 1987, total precipitation during these 4 months was 218.8
mm. but flushing peaked later that year.
The spring application of fertilizer was applied in February 1985, March 1986 and
February 1987. The timing of the spring flush was apparently unrelated to the timing
of fertilizer applications.

Howard & Solis: Mahogany Webworm 479

From the above observations it appears that daylength and temperature are impor-
tant in timing the flight of mahogany webworm moths, the spring flush of West Indies
mahogany foliage, and the occurrence of mahogany webworm populations. These re-
lationships are probably subtle and complicated and remain to be proven.


We wish to thank J. V. DeFilippis for excellent technical assistance and for Figures
1 & 2. N. E. Woodley, P. M. Marsh (Systematic Entomol. Lab., IIBII, SEA, U.S.
Department of Agriculture, Beltsville, MD.), and B. E. Cooper (Biosystematic Res.
Cent., Agriculture Canada, Ottawa) kindly identified specimens of parasitoids. We
thank T. K. Broschat and R. H. Scheffrahn, University of Florida, for reviewing the
manuscript. This is Florida Agricultural Experiment Station Journal Series No. 9575.


ALVIM, P. de T. 1964. Tree growth periodicity in tropical climates, pp. 479-495 In M.
H. Zimmermann [ed.], Formation of wood in forest trees. Academic Press, New
BROSCHAT, T. K., AND H. M. DONSELMAN 1983. Effect of photoperiod on growth
of West Indian mahogany. HortScience 18: 206-207.
CHELLMAN, C. W. 1978. Pests and problems of south Florida trees and palms. Florida
Dept. Agric. & Consum. Serv., Div. Forestry, Tallahassee, Fl. 103 pp.
LAMB, F. B. 1966. Mahogany of tropical America; Its ecology and management. Uni-
versity of Michigan Press, Ann Arbor. 220 pp.
LONGMAN, K. A., AND J. JENIK. 1974. Tropical forest and its environment. Longman
Group Ltd., London. 196 pp.
Comparisons between blacklight and pheromone traps for monitoring the west-
ern bean cutworm (Lepidoptera: Noctuidae) in south central Idaho. J. Econ.
Entomol. 80: 242-247.
MORTON, J. G. 1987. Our misunderstood mahogany and its problems. Proc. Florida
State Hort. Soc. 100: 189-195.
PENNINGTON, T. D., B. T. STYLES, AND D. A. H. TAYLOR. 1981. Meliaceae. Flora
Neotropica Monograph No. 28. New York Bot. Gard., Bronx, N.Y.
PETERSON, A. 1963. Egg types among moths of the Pyralidae and Phycitidae -
Lepidoptera. Florida Entomol. Suppl. No. 1.: 1-14.
REINERT, J. A., AND F. W. HOWARD. 1982. Susceptibility of the mahogany web-
worm to insecticides. Proc. Florida State Hortic. Soc. 95: 288-289.

~ .4


em *

;45 &

Florida Entomologist 72(3)


'Fort Lauderdale Research and Education Center,
University of Florida
Institute of Food and Agricultural Sciences
Fort Lauderdale, Florida 33314
2Coconut Research Division
Ministry of Food Production,
Marine Exploitation, Forestry, and the
Environment of the Republic of Trinidad and Tobago, West Indies


Larvae of Rhynchophorus cruentatus (F.) of variable starting weights gained weight
when incubated at 23 C for 21 days in sugarcane stems. R. cruentatus was reared
through one generation on sugarcane stems with a mean generation time of > 212 days
at 23 C. Comparisons were made with the syncarpium of pineapple as a growth medium
for R. cruentatus at 23 C and 28 C for 25 days. Mean fecundity of R. cruentatus on
pineapple was not significantly different at the two temperatures or between newly-
emerged field-collected adults or those reared on sugarcane. Larvae from the pineapple
syncarps at 280 C were significantly heavier than those reared at 230 C for 25 days.
Complete development of R. cruentatus did not occur on pineapple. From these prelim-
inary studies a method for laboratory rearing R. cruentatus was developed. Single pairs
of newly-emerged females and males were confined to individual pineapple syncarps for
> 25 days at 280 C. Resultant larvae were individually transferred into sugarcane stems
to complete metamorphosis. Using a pineapple-sugarcane diet, R. cruentatus was
reared through five successive generations in Florida and R. palmarum (L.) was reared
through one generation in Trinidad.


Larvas de Rhynchophorus cuentatus (F.) de various pesos, al principio aumentaron
en peso cuando fueron incubadas a 230 C por 21 dias en cafias de azucar. Rhynchophorus
cruentatus fue criado por una generaci6n en cafias de azucar con un tiempo promedio
de generaci6n de > 212 dias a 230 C. Se hicieron comparaciones con el sincarpo de piia
como un medio para criar R. cruentatus a 23 C y 280 C por 25 dias. La fecundidad
promedio de R. cruentatus en pifia no fue significantemente diferente a las dos tem-
peraturas, entire adults criados en cafa de azucar, ni entire los colectados en el campo.
Las larvas criadas en el sincarpo de pifia a 280 C eran significantemente mAs pesadas
que las criadas a 230 C por 25 dias. No. hubo metam6rfosis complete de R. cruenttus en
la pifia. De estos studios preliminares, se desarroll6 un m6todo para criar R. cruentatus
en el laboratorio, en el cual se encerraron pares de hembras y machos recientemente
emergidos en sincarpos individuals por > 25 dias a 280 C. Larvas resultantes se tras-
ladaron individualmente a cafias de azucar para completar la metam6rfosis. Sobre una
dieta de pifia-cafia de azfcar, se cri6 R. cruentatus por 5 generaciones sucesivas en la
Florida, y se cri6 R. palmarum por una generaci6n en Trinidad.

Rhynchophorus cruentatus (F.) is a pest of transplanted cabbage palmettos, Sabal
palmetto (Walter), and Canary Island date palms, Phoenix canariensis Hortorum ex
Chabaud, in Florida (Giblin-Davis & Howard 1989). These palms are commonly used as

September, 1989

Giblin-Davis et al.: Rearing Palm-infesting Weevils

woody ornamentals in urban landscapes in Florida. Most species of Rhynchophorus are
pests of economically important species (e.g., coconut palm, Cocos nucifera L., and oil
palm, Elaeis guineensis Jacquin) (Wattanapongsiri 1966, Giblin-Davis & Howard 1989).
The palm weevil, R. palmarum (L.) is unique in the genus because it vectors the red
ring nematode, Rhadinaphelenchus cocophilus (Cobb), to apparently healthy palms in
the southern Caribbean, Mexico, and Central and South America (Griffith 1987). The
red ring nematode causes the red ring disease of coconut and oil palms and is usually
manifested as a lethal wilt of 3-10 year old hosts (Griffith 1987). The association between
the red ring nematode and R. palmarum is presumed to be recent in origin and other
species of Rhynchophorus may be potential vectors (Griffith 1987). R. cruentatus is
distributed in the southeastern United States (Wattanapongsiri 1966, Woodruff 1967)
and known to attack Phoenix palms which are hosts of the red ring nematode in the
Neotropics (Esser 1969, Giblin-Davis & Howard 1989).
Research on the red ring nematode vector potential, biology, and control of R.
cruentatus requires an effective method of laboratory rearing. Presently, no culture
method exists for R. cruentatus so that all work has been done with individuals collected
from the field (Giblin et al. 1987, Giblin-Davis & Howard 1989). Berger (1907) reared
two field-collected larvae of R. cruentatus to adults on buds of the saw palmetto, Ser-
renoa repens (Bartram). Rhynchophorus ferrugineus Oliver and R. palmarum have
been reared in the laboratory on cut petiole or stem tissue of coconut palms, but this
was expensive and time consuming (Rananavare et al. 1975, Wilson 1963). Sugarcane,
Saccharum officinarum L., was reported to be a good substitute for coconut stem for
laboratory rearing of R. ferrugineus (Rahalkar et al. 1972). The method for rearing R.
ferrugineus was improved by using shredded sugarcane stem tissue for oviposition,
sugarcane shreds in nutrient agar for newly hatched larvae, and sugarcane stem pieces
for older larvae (Rananavare et al. 1975). Both R. palmarum and R. cruentatus have
been reported from pineapple, Anana comosus (L.), (Wattanapongsiri 1966) but no
reports exist on whether either can complete its life cycle on this host.
The objectives of our study were: 1) to evaluate sugarcane stem and the syncarpium
of pineapple for laboratory growth and maintenance of R. cruentatus; 2) to optimize a
rearing procedure for R. cruentatus and test it on R. palmarum and 3) to assess the
longevity and fecundity of newly-emerged field-collected R. cruentatus under labora-
tory conditions.


Laboratory Growth. Larvae of R. cruentatus were collected from mature infested
cabbage palmettos from Broward and Collier counties, FL, as described by Giblin-Davis
and Howard (1989). Each larva was washed in tap water, patted dry, weighed, and
placed into a 1.1 by 5.0 cm hole drilled into a 40-45 cm length of sugarcane stem (300-500
g) (variety Yellow Gal or 1527). Canes were confined individually in clean, plastic soda
bottles (2 liter), with tops removed. Bottles were kept in large garbage cans tightly
covered with fine mesh nylon screen covers (300 Jm openings). The canes were har-
vested after 21 or 42 days at 230 2 C and the stage, weight, and appearance of the
insect recorded. Experiments were replicated twice with at least 12 larvae per replicate.
Larvae from the first 21-day harvest were transferred into sugarcane as described
above for another 21-day period to assess subsequent growth. Weight for each larva
after 21 or 42 days was compared with its starting weight.
A separate experiment was done as described above with progeny from field-col-
lected adults of R. cruentatus that were allowed to mate and oviposit in groups of 10
20 on 10 cm pieces of split sugarcane stem for 1 week. Larvae were harvested 32 days
later and subcultured successively on new pieces of sugarcane stem every 21 days in
an attempt to rear them from egg to adult. Larvae were transferred five times into

482 Florida Entomologist 72(3) September, 1989

sugarcane stems 40-45 cm long. At this point all surviving larvae were > 3.0 g and were
transferred into 23-25 cm lengths (170-282 g) without subsequent transfer. There were
five replicates in time. Changes in larval weight after 21 days in sugarcane stem were
compared with the starting weight.
Pineapple syncarpium, the fleshy collective fruit of the pineapple, was tested for its
suitablity for rearing R. cruentatus at 23 2 C or 28 20 C for 25 days. Pupae of R.
cruentatus in cocoons from several infested mature cabbage palmettos collected from
Collier Co., FL, on 4 May, 1987 were placed in 100 ml plastic containers with a mois-
tened Kimwipe tissue paper (Kimberly-Clark Corp., Roswell, GA) with 4 ml deionized
water per container at 280 C. Newly-emerged adults were collected daily, weighed, and
used for experiments. Newly-emerged adults that had been reared through one gener-
ation on sugarcane were also used in these tests. One male and one female weevil, was
confined on a fresh pineapple syncarp (1100-1500 g) in clean polyethylene buckets (19
liter) with fine-screened tops (300 IJm openings). After 25 days, the pineapple syncarp
was harvested and the eggs and larvae were counted and weighed. Comparisons of
fecundity and larval weight were made with PROC GLM and separation of means was
done with the Waller Duncan option (P = 0.05); correlations were done with PROC
CORR (SAS Institute 1985). Three replicates with at least three syncarps per replicate
were done, except for adults cultured in sugarcane (two replicates).
Rearing Optimization. Results from the preceding section were used to optimize
rearing of R. cruentatus. A reproductive pair of newly-emerged weevils was confined
to the fresh syncarp of a pineapple (1100-1500 g) in a screened container (19 liter) as
described above for 25 days at 28 2 C. Larvae from the pineapple were harvested,
washed, patted dry, weighed, and placed into holes drilled into 23-25 cm lengths of
sugarcane stem (170-282 g) and incubated at 28 2 C for > 21 days. Beginning at 22
days, canes were split open lengthwise to examine the status of the insect. Dead insects
were discarded; those that had pupated were transferred in their cocoon to a vented
container (100 ml) with a moistened tissue as above. Weevils were monitored regularly
for emergence, at which time they were weighed and used to rear successive genera-
tions or to assess longevity for cultured weevils (see next section). Generation time was
calculated as the time from adult confinement on pineapple to emergence of adults of
the next generation, unless otherwise stated. Comparisons for sex-related differences
in larval weight at pineapple harvest, generation time, and emergence time from pineap-
ple harvest were made with a PROC GLM, and correlations were done with PROC
CORR (SAS Institute 1985).
The pineapple-sugarcane rearing method was attempted on R. palmarum at the
Central Experiment Station, Centeno, Trinidad. All experiments in Trinidad were done
with locally produced pineapples (about 500-600 g) and sugarcane stems (about 50 cm
Longevity and Fecundity. Longevity and fecundity of R. cruentatus was assessed
with field-collected pupae which were harvested and allowed to emerge as adults as
described above. To determine longevity without food, a newly-emerged male and
female of R. cruentatus were confined to a vented plastic container (100 ml) with a
moistened Kimwipe. The tissue was changed and moistened with 4 ml of deionized
water once per week. To assess longevity and fecundity with food, weevils were con-
fined in pairs as above in 100 ml plastic containers with a 35-40 g slice of fresh pineapple.
The pineapple slice was replaced weekly and the old piece was confined to a separate
container for 7 days before dissection and examination for weevil eggs and/or larvae.
Weevils were examined daily for mortality. Adults that were still alive at 63 days were
weighed for comparison with their starting weight. There were four replicates in time
with five pairs of weevils per replicate for each of the moistened tissue and pineapple
longevity assays. Data were analyzed as described previously.

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