(An International Journal for the Americas)
Volume 69, No. 1 March, 1986
TABLE OF CONTENTS
Announcement 69th annual meeting and call for papers ................................ i
THE FLORIDA ENOMOLOGICAL SOCIETY'S FIRST CARIBBEAN CONFERENCE
OF E NTOM OLOGY ......................................................... ................ ......... 1
POPENOE, H.-The International Dimensions of Florida Agriculture: A Syn-
opsis ...................................... ............. .......... ............ ................. 3
BRAITHWAITE, C. W. D.-IICA Plant Protection Programme in the Caribbean
A Case of Technical Cooperation in Agricultural Development Assist-
ance ......... ... ... ................................................ ...... 5
SHANKLAND, D. L., AND H. N. NIGG-An Environmental Toxicology Program
for F lorida .................................................................................... 15
HOLLIS, W. L.-Cooperative Industry Efforts with Developing Countries to Im-
prove Agrochemical Registration, Labeling and Education and Training
Program s ....................................................... ........ 21
KING, A. H,.-Latin American Entomological Serials .................................. 30
SYMPOSIUM: INSECT BEHAVIORAL ECOLOGY-85
WALKER, T. J.-Stochastic Polyphenism: Coping With Uncertainty ............... 46
RAUSHER, M. D.-Competition, Frequency-Dependent Selection, and Diap-
pause in Battus philenor Butterflies .................................... ....... 63
MCNEIL, J. N.-Calling Behavior: Can It Be Used to Identify Migratory Species
of M oths? ...................................................................................... . ......... 78
QUINN, J. S., AND S. K. SAKALUK-Prezygotic Male Reproductive Effort in
Insects: Why do Males Provide More Than Sperm? ............................ 84
PIERCE, J. D., JR.-A Review of Tool Use in Insects .................................. 95
MORIN, J. G.-"Firefleas" of the Sea: Luminescent Signaling in Marine Ostra-
code Crustaceans ........................................ .......... .................... 105
TURNER, M. E.-Multiple Mating, Sperm Competition and the Fertility Com-
ponent of Fitness in Drosophila pseudoobscura .................................. 121
LLOYD, J. E.-Behavior Ecology Milestones ..................... ................ 129
SYMPOSIUM: USE OF PHEROMONES IN TROPICAL CROPS
M ITCHELL, E. R.- Preface .................................................................. 131
MITCHELL, E. R.-Pheromones: As the Glamour and Glitter Fade-The Real
W ork Begins ........................................................ ...................... 132
SILVAIN, J. F.-Use of Pheromone Traps as a Warning System Against Attacks
of Spodoptera frugiperda Larvae in French Guiana ......................... 139
DICKERSON, W. A.-Grandlure: Use in Boll Weevil Control and Eradication
Programs in the United States .................................................. .. 147
WHITCOMB, W. H., AND R. M. MARENGO-Use of Pheromones in Boll Weevil
Detection and Control Program in Paraguay ..................................... 153
SIVINSKI, J. M., AND C. 0. CALKINS-Pheromones and Parapheromones in the
Control of Tephritids ............................. .. .... ............... 157
Continued on Back Cover
Published by The Florida Entomological Society
FLORIDA ENTOMOLOGICAL SOCIETY
OFFICERS FOR 1985-86
President .................................... ................. D. H. Habeck
President-Elect ................................. ....................... D. J. Schuster
Vice-President ........................................ ................ J. L. Taylor
Secretary ................................................... E. R. Mitchell
Treasurer ........................................................ A. C. Knapp
S M. L. Wright, Jr.
J. E. Eger, Jr.
R. C. Bullock
Other Members of the Executive Committee ....... Bullock
C, G. Witherington
J. R. McLaughlin
Editor .................................. ................. J. R. McLaughlin
Associate Editors ...................................................... ....................... A. Ali
C. S. Barfield
J. B. Heppner
M. D. Hubbard
0. Sosa, Jr.
H. V. Weems, Jr.
W. W. Wirth
Business Manager ....................................... ..... .................. A. C. Knapp
FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and De-
cember. Subscription price to non-members is $30.00 per year in advance, $7.50 per
copy. Membership in the Florida Entomological Society, including subscription to Flor-
ida Entomologist, is $25 per year for regular membership and $10 per year for students.
Inquires regarding membership, subscriptions, and page charges should be addressed
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.
Authors should consult "Instructions to Authors" on the inside cover of all recent
issues while preparing manuscripts or notes. When submitting a paper or note to the
Editor, please send the original manuscript, original figures and tables, and 3 copies
of the entire paper. Include an abstract and title in Spanish, if possible. Upon receipt,
manuscripts and notes are acknowledged by the Editor and assigned to an appropriate
Associate Editor who will make every effort to recruit peer reviewers not employed by
the same agency or institution as the authors(s). Reviews from individuals working
out-of-state or in nearby countries (e.g. Canada, Mexico, and others) will be obtained
where possible. Page charges are assessed for printed articles.
Manuscripts and other editorial matter should be sent to the Editor, JOHN R.
MCLAUGHLIN, 4628 NW 40th Street, Gainesville, FL 32606.
This issue mailed May 23, 1986
THE 69th ANNUAL MEETING OF THE
FLORIDA ENTOMOLOGICAL SOCIETY
ANNOUNCEMENT AND CALL FOR PAPERS
The J4wida Entomological Society will hold its 69th Annual Meeting on
6-8 Augitf~1986 at the Sheraton Sand Key Hotel, Clearwater Beach, Florida. The
location is 1160 Gulf Boulevard, Clearwater Beach, Florida 33515; telephone-1-813-595-
1611. Room rates will be $66.00 either single or double. Pre-registration and registration
fees will be released in the June, 1986 Florida Entomologist and the April Newsletter.
Since many will present papers please copy the sheet and submit before 1 June 1986,
JAMES L. TAYLOR, Chairman
Program Committee, FES
P. O. Box 1893
Sanford, Florida 32771
Eight minutes will be allotted for presentation of oral papers, with 2 minutes for
discussion. In addition, there will be a separate session for members who may elect to
present a Project (or Poster) Exhibit. The three oral student papers judged to be the
best on content and delivery will be awarded monetary prizes during the meeting.
Student authors must be Florida Entomological Society Members and must be regis-
tered for the meeting.
S rM a
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H a, e
^ ?S ^
u o s'
THE FLORIDA ENTOMOLOGICAL SOCIETY'S
FIRST CARIBBEAN CONFERENCE OF ENTOMOLOGY
August 5-8, 1985
Ocho Rios, Jamaica, West Indies
Some came to enjoy the splendor of Jamaica, others to share entomological informa-
tion and discuss common interests in insect biology and control, but all were treated to
an incredible potpourri of Caribbean agriculture. We advisedly selected Ocho Rios,
Jamaica, as the site for the first meeting because of the Jamaicans' desire to participate,
the relevance of their agriculture to other areas of the Caribbean, the relative political
neutrality and security of the country, and the availability of required logistical services.
Aesthetic appeal also was important, along with a diversity of associated activities such
as field trips to see crops and their associated harmful and beneficial insects, natural
sites for collecting insects, and existing entomological collections and expertise. Our
ambitious objectives for this pioneering meeting were to strengthen professional re-
lationships with colleagues in entomology and plant protection, foster collaboration
among Caribbean, Latin American, and U. S. scientists, encourage the exchange of
scientific information on tropical and subtropical entomology as it impinges on agricul-
tural production and product protection, expose practitioners to the technical problems
and singular opportunities associated with developing tropical agriculture, and ulti-
mately evaluate the Caribbean Conference and determine if others should be held.
From the onset, we felt that the potential pay-offs of such a meeting could be enormous.
We were welcomed to the island on Tuesday morning, August 5, by Dr. Osmond
Tomlinson, Custos of St. Ann Parish including Ocho Rios, who piqued our curiosity
about current and historical associations among countries that lie within or border the
Caribbean. This provocative introduction, the keynote address, and four invitational
papers preceded a symposium on the "Biology and Control of Coffee Pests" organized
and presided over by Ms. Janice Reid of Jamaica. Topics included the coffee berry
borer, other pests of coffee, and both chemical and biological control. Submitted papers
on crop protection moderated by Dr. Gary Leibee followed describing the coffee leaf-
miner in Puerto Rico; pink bollworm in Peru; mites in Central America and the West
Indies; the potato tubermoth in Venezuela; Liriomyza, weevils, fruit flies, and insec-
ticide testing in Florida; and cultural and chemical control of bean slug in Honduras.
The afternoon was consumed by concurrent sessions with Dr. Russ Mizell's urban en-
tomology and Dr. Emil Moherek's crop protection papers opposite those on citrus and
medical entomology moderated by Dr. Bob Bullock and Dr. Arshad Ali, respectively.
The exceptionally wide range of subjects were presented by speakers from Guadeloupe,
Canada, Jamaica, Cayman Islands, and the U.S. The traditional "Behavioral Ecology"
symposium produced by Dr. Jim Lloyd and "Student Paper Contest" officiated by Dr.
Nan-yo Su on Wednesday were accompanied by poster exhibits and Dr. Everett Mitch-
ell's symposium on the "Use of Pheromones in Tropical Crops". Thursday morning was
devoted to "Biological Control" under Dr. Fred Bennett and "Crop Protection in Small
Farm Systems" with Dr. Keith Andrews. Of the 98 non-student papers listed in the
program, 33 were from other than U.S. with participation by a total of 14 countries.
One of the highlights of the meeting was a Thursday afternoon and eveniug "Citrus
Tour" organized by Dr. Joe Knapp and conducted by Dr. Villi Bent. We traveled south
through the central highlands of Middlesex to groves near Linstead. There we gathered
in the field to meet Mr. Walter van Whervin, the Ministry of Agriculture's Chief of
Plant Protection. Others present from Jamaica included Mr. David McConnell (Chair-
Florida Entomologist 69(1)
man, United Estates Limited), Mr. Mark McConnell (Managing Director, United Es-
tates Limited), Mr. Robert Clarke (Manager, Worthy Park Farms), Mr. Winston Miller
(Manager, Winston A. B. Miller, Ltd.), Mr. Ken Newman (Managing Director,
Wakefield Farms, Ltd.), Mr. S. S. Stons (Project Manager, Citrus Rehabilitation
Scheme), Mr. Crook (Expatriate Advisor), Mr. L. A. Bell (Extension & Research Of-
ficer), Mr. Amir (Expatriate Advisor, World Bank Fund), Mr. W. E. Tavares (Field
Manager, United Estates, Ltd.), and Mr. L. V. Bowie (Ass't. Field Manager, United
Estates, Ltd.). After this brief but impressive ceremony, the owners (David, Peter,
Stuart and Mark McConnell) showed us fields of orange trees, sugarcane, and coconut
palms. Among other things, they explained cultural practices and experiments designed
to control the citrus root weevil. Our next stop was a nursery for citrus and tropical
fruit, such as guava and passionfruit, that supplied the plants for new groves. Finally,
we entered the St. Thomas-Ye-Vale Valley that had such a severe infestation of citrus
root weevils that we were able to collect specimens merely by walking around the trees.
On the way home, to our surprise and delight, our hosts treated us to a veritable feast
of delicate meat pies and pastry served with hand-squeezed orange and tropical fruit
juices. The tour was enjoyed and appreciated by all as a special experience in agricul-
ture, science, natural history, and Jamaican hospitality.
At the traditional "Bull Session", we had an opportunity to ask those in attendance
to fill out a brief questionnaire and carry on a rather lengthy discussion about how they
would evaluate this first international venture. In summary, the group responding
there (and later) voiced overwhelming support for the Florida Entomological Society to
continue to hold periodic meetings in the Caribbean Basin. Instead of every 5 years,
most felt that a 3 year schedule was even better.
It occurred to us that most of the stated objectives of the Caribbean Conference
were accomplished and that this success should be documented. Therefore, we assem-
bled the keynote address, opening invitational papers and a listing of "Latin American
Entomological Serials" into a single unit for publication in Florida Entomologist. Dr.
Hugh Poponoe put in perspective the commonality of agriculture in countries that share
the Caribbean, Dr. Chelston Brathwaite identified the primary agricultural institutions
of the Caribbean and briefly discussed the functions of each, efforts to effectively
monitor and control hazardous chemical waste in Florida, a relevant problem for other
areas of the Caribbean, were described by Drs. Dan Shankland & Herb Nigg, and an
international "crisis" in pesticide development, production, and regulation in developing
countries was exposed by Dr. Bill Hollis. The serials list was compiled by Ms. Ann
King. We sincerely thank all the participants for their contributions to this first interna-
tional Caribbean Conference on Entomology.
C. S. Barfield and N. C. Leppla
Local Arrangements Committee
Caribbean Conference 3
THE INTERNATIONAL DIMENSIONS OF
H. L. POPENOE
Director for the Center for Tropical Agriculture
University of Florida
Institute of Food and Agricultural Sciences
Gainesville, Florida 32611 USA
There are very close bonds that exist among agricultural scientists and practitioners
in the Caribbean Basin. Florida shares a common ecology and common crops with
Caribban nations to a greater degree than with most states in the contiguous United
States. Thus, continuing and emerging entomological problems in Florida require shar-
ing of information and cooperative research efforts with other members of the Caribbean
Existen estrechos vinculos entire los cientificos agricolas y practicantes en el Area
del Caribe. La Florida compare una ecologia y cultivos en comfim con naciones del
Caribe en un mayor grado que con la mayoria de los estados contiguos de los Estados
Unidos. De aqui que continues y salientes problems entomol6gicos en la Florida re-
quieren el intercambio de informaci6n y esfuerzos cooperatives de investigaci6n con
otros miembros de la comunidad del Caribe.
We truly live in an interdependent world. In entomology, perhaps more than any
other science related to agriculture, both our problems and their solutions are often to
be found in some other country. This is especially true throughout the Caribbean Basin
where a large number of tropical and sub-tropical countries are in close proximity. The
United States is clearly represented in the region through the state of Florida. This
explains, to a great extent, Florida's affinity to the Caribbean and underscores the
importance of international programs (Fig. 1).
Florida is a peninsula surrounded by water. It doesn't share common ecology or
common crops with other states. We actually have more in common agriculturally with
nations to the south than with most states in the contiguous United States. Thus, since
Florida shares agricultural and urban problems related to entomology with virtually
every country in the Caribbean, information is shared across international boundaries.
An opportunity exists for us to conduct cooperative research on incipient pests,
those that have been controlled or eradicated but remain a constant threat, and even
some that are currently of unknown importance. Hopefully, these activities will continue
to serve as an early warning system and prepare researchers to deal with damaging
insects that are detected within our borders. The international dimension of this work
allows us to be prepared for the current and potential realities of introduced agricultural
In Florida agriculture, entomologists have been repeatedly called on to solve novel
problems, in record time, to save growers, producers and consumers estimated millions
of dollars. This phenomenon has become an expectation rather than an extraordinary
feat. Through the cooperative efforts of our international programs we are often able
Florida Entomologist 69(1)
Fig. 1. Caribbean Basin Map emphasizing the proximity of Florida. (Source un-
to anticipate problems before they emerge and predict the outcome of available insect
control practices. This shared knowledge and the collaboration of international scientists
has proven its worth time and again.
Recent examples of continuing or emerging problems and allied foreign research
programs include work on medflies, citrus blackflies, mole crickets, fire ants, sugarcane
rust, citrus canker, and lethal yellowing of coconuts. We also have cooperative projects
on insect-related livestock diseases; African swine fever, heartwater disease, blue
tongue, and equine encephalitis have been of particular importance.
International programs also give us an opportunity to better manage our crops,
develop new commodities, and open or expand markets. Increasingly populous and
cosmopolitan societies of the Caribbean community (including Florida) demand a greater
quantity, quality, and variety of food and fiber. Therefore, we have established coopera-
tive efforts on various crops including winged bean, malanga, carambola, sugarcane and
Specifically, the Government of Jamaica and the University of Florida have had
cooperative projects in such diverse areas of mutual interest as: agricultural mechaniza-
tion and dairy cooperatives with the Ministry of Agriculture; the Jamaica School of
Agriculture through the Ministry of Education; Lethal Yellowing of coconut in conjunc-
tion with the Coconut Board; and, more recently, the Farming Systems Support Project
and agricultural marketing with EXPO 21. We have also had individual scientists work-
ing on muck soils, sugarcane, root crops, aquaculture, and parasites to control insects
The Florida Entomological Society is to be commended for its decision to hold its
68th Annual Meeting in Jamaica, emphasizing the international dimension of its ac-
tivities. It is a fitting recognition and a tribute to the very close bonds that exist among
agricultural scientists and practitioners in the Caribbean Basin. Hopefully this is a
precedent for future opportunities for the Society to be hosted by other countries in the
region, to exchange information, promote friendship and further collegial relations.
Caribbean Conference 5
IICA PLANT PROTECTION PROGRAMME
IN THE CARIBBEAN
A CASE OF TECHNICAL COOPERATION IN
AGRICULTURAL DEVELOPMENT ASSISTANCE
CHELSTON W. D. BRATHWAITE
Regional Plant Protection Specialist
IICA Office in Trinidad and Tobago
The IICA Plant Protection Programme for the Caribbean seeks to promote and
support the efforts of the countries of the Caribbean to prevent and reduce crop losses
caused by pests, diseases and weeds. The programme, which was started in 1981, has
as a fundamental strategy reciprocal technical cooperation where the experiences and
technical information of some countries are used in a transfer of technology to others
utilising human resources, information exchange and mechanisms for communication.
The establishment of a Society for Plant Protection in the Caribbean, the establishment
of a Regional Plant Protection Newsletter and annual meetings of Heads of Plant Pro-
tection represent the major mechanisms in this technical cooperation package.
Attempts to harmonize pesticide legislation and the training and certification of
Plant Quarantine Inspectors represent approaches to standardize the legislative aspects
of Plant Protection in the region.
Initiatives have also been focused on survey and eradication studies and proposals
are imminent to set up a data base of Plant Protection information for the Caribbean.
El Program de Protecci6n de Plantas IICA para el Caribe busca promover y apoyar
el esfuerzo de los paises del Caribe para prevenir y reducir p6rdidas, en los cultivos
causadas por plagaas, enfermedades y malezas. El program que comenz6 en 1981, tiene
como estrategia fundamental la reciprocidad de cooperaciofi t6cnica donde la experiencia
e informaciofi tecnica de algunos paises son usadas en una transferencia de tecnologA a
otros utilizando fuentes humans, intercambio de informaci6n, y mecanismos para la
comunicaci6n. El establecimiento de una Sociedad para la Proteccion de Plantas en el
Caribe, el establecimiento de un Boletin Regional para la Protecci6n de Plantas, y una
reunion annual de los Directores de Protecci6n de Plantas, representan los principles
mecanismos en este paquete de cooperaci6n tCnica.
Intentos para harmonizar legislaci6n sobre pesticides y el entrenamiento y certifica-
ci6n de Inspectores de Cuarentena de Plantas, representan acercamientos para hacer
uniform los aspects de Protecci6n de Plantas en la region.
Tabien se han concentrado iniciativas sobre studios de encuesta y erradicaci6n, y
proposiciones para establecer una base de datos de informaci6n sobre Protecci6n. de
Plantas para el Caribe son iminentes.
WHAT IS IICA
The Inter-American Institute for Cooperation on Agriculture-IICA-is an interna-
tional, inter-governmental organization specialized in agriculture. It is governed by its
own Convention and has been recognized as a specialized Inter-American Agency under
the Charter of the Organization of American States.
The purposes of IICA are to "encourage, promote, and support the efforts of the
Member States to achieve their agricultural development and rural well-being".
Florida Entomologist 69(1)
The Institute was founded in 1942 as the Inter-American Institute of Agricultural
Sciences. On December 8, 1980, a new Convention was ratified. Under this new Conven-
tion, the Institute changed its name to the Inter-American Institute for Cooperation on
Agriculture, expanded its purposes and altered its institutional structure.
IICA has 29 Member States: Argentina, Barbados, Bolivia, Brazil, Canada, Chile,
Colombia, Costa Rica, Dominica, Dominican Republic, Ecuador, El Salvador, Grenada,
Guatemala, Guyana, Haiti, Honduras, Jamaica, Mexico, Nicaragua, Panama, Paraguay,
Peru, Saint Lucia, Suriname, Trinidad and Tobago, United States of America, Uruguay
and Venezuela. Twelve Observer Countries contribute to Institute activities: Austria,
Belgium, Egypt, France, Germany, Israel, Italy, Japan, Korea, the Netherlands, Por-
tugal and Spain. IICA has a technical staff of 180 international professionals. Around
1,200 persons are working for the Institute throughout the hemisphere.
IICA's resources flow from annual quotas which the member countries commit them-
selves to pay each year. Funds also derive from agreements, contracts, contributions,
and grants for which the Institute signs with other national and international organiza-
tions. For 1984, the Institute's highest governing body, the Inter-American Board of
Agriculture, has approved a budget of 37 million dollars.
IICA's Director General is Venezuelan scientist and educator, Dr. Francisco Morillo
Andrade. The Deputy Director is Dr. Quentin M. West of the United States.
WHAT IICA DOES
IICA concentrates its action in ten hemisphere-wide programs, which provide a
framework for the annual performance of over a thousand activities. These activities
are carried out through agreements reached with the Governments of the Member
States, and are in the hands of decentralized technical teams covering the 29 Member
IICA's hemisphere-wide programs are: Formal agricultural education; Support of
national institutions for the generation and transfer of agricultural technology; Conser-
vation and management of renewable natural resources; Animal health; Plant protec-
tion; Stimulus for agricultural and forest production; Agricultural marketing and agro-
industry; Integrated rural development; Planning and management for agricultural de-
velopment and rural well-being; and Information for agricultural development and rural
The specific objectives over the medium term have been defined in accordance with
the general objectives and overall strategy of IICA, and on the basis of the concepts of
agricultural development and rural well-being that are put forth in the guidelines for
the Institute's general policies. These specific objectives include cooperating with the
Member States through:
a. Bringing about the growing, effective participation of rural dwellers, especially
the low-income strata, in decision-making on projects affecting them, seeking to incor-
porate them fully into the benefits of economic and social progress.
b. Developing human resources by promoting formal and non-formal training, to
improve productive efficiency and promote the participation of the rural population in
processes for achieving rural well-being.
c. Developing and consolidating national systems for the generation and transfer of
technology, in order to help each country fit itself into the regional and world technolog-
ical framework. This would be done for the purpose of improving both agricultural and
forest production and productivity, preventing and reducing losses to pests and diseases
in crops and herds, and maximizing the use and conservation of renewable natural
Caribbean Conference 7
d. Developing policies, mechanisms and tools for stimulating the efficient production
and marketing of inputs and of agricultural, livestock and forest products, domestically
e. Reinforcing regional and integrated rural development institutions for planning
and implementing integrated projects, so as to coordinate institutional action and pro-
vide for the effective participation of beneficiaries.
f. Reinforcing public and private institutional systems in the many facets of setting
national goals, planning, and implementation at all levels, on the basis of the retrieval
and analysis of information for better defining and implementing policies and programs
of agricultural development and rural well-being, and for establishing IICA's own
priorities for action.
THE PLANT PROTECTION PROGRAMME
The Ministers of Agriculture attending the VII Inter-American Conference of Agri-
culture held in Honduras in 1977 expressed their concern regarding the disease prob-
lems of plants and animals throughout the Western Hemisphere. Two of the ten recom-
mendations made at this conference refer to this subject. The Special Committee of the
Eighteenth Annual Meeting of IICA Board of Directors held in October, 1978 recom-
mended that the Director General of IICA study a proposal aimed at the establishment
of a mechanism for the coordination of efforts to fight pests and disease problems
affecting animals and plants and which are the cause of significant losses in the Hemis-
As a result of these directives, IICA has established a Hemispheric Plant Protection
Programme designed to prevent, control and if possible, eradicate diseases and pests
which cause economic damage to crops in the Hemisphere and which threaten to spread
to other regions. The programme is made up of a Programme Director stationed at
IICA headquarters in San Jose, Costa Rica and four Plant Protection Specialists, one
stationed in each of the four regions of the Hemisphere. The Plant Protection Specialist
for the Caribbean is Chelston W. D. Brathwaite, Plant Pathologist stationed in the
IICA Office in Trinidad and Tobago.
In accordance with IICA's basic strategy, this programme is directed towards
strengthening national and regional efforts being carried out by other organizations. It
is designed to support, coordinate and collaborate with other International, regional
and subregional institutions working in this area and in no case will duplicate or replace
The programme recognizes that the spread of pests, diseases, and weeds that affect
basic food and export crops aggravate the food, foreign exchange and energy needs of
the Latin American and Caribbean countries. Coordinated international action can con-
tribute to reducing the spreading and incidence of these pests, weeds and diseases,
since the individual capabilities of national plant protection institutions are usually li-
mited by low levels of physical, human and financial resources with which to attain their
General Objective of the Programme-To promote and support the efforts of the coun-
tries to prevent and reduce crop losses caused by pests, diseases and weeds.
Specific Objectives of the Programme-To cooperate with the countries in expanding
and improving their institutional capability to:
a. Update and standardize national and international legal provisions and regulations
governing plant protection.
b. Identify, detect and estimate the damage caused by the main crops pests, diseases
c. Plan, coordinate and implement programs for reducing the incidence and prevent-
Florida Entomologist 69(1)
ing the spread of the main crop pests, diseases and weeds.
d. Plan, coordinate and implement research and technical exchange programs on
crop pests, diseases and weeds.
e. Generate mechanisms for upgrading the physical, human and financial resources
of plant protect institutions, according to the levels of responsibility that have been
Strategy of the Programme-To promote and support:
a. The updating and standardization of national and international legal provisions
and regulations governing plant protection (quarantine and pesticides).
b. The formulation, implementation and evaluation of multinational projects that
involve economically important pests and diseases of mutual interest to several coun-
c. The formulation, implementation and evaluation of high-priority projects at the
d. The use of technical and human resources from other IICA programs, from
CATIE, and from national and international institutions with experience in this field.
e. The operational and technical reinforcement of national and international institu-
tions working in this field (OIRSA, FAO, CIP, NAPPO, CIAT, CIMMYT).
f. Coordination with other international agencies.
g. The organization and promotion of meetings, seminars and other events for con-
sultation and orientation to establish working guidelines and priorities for action.
h. The organization of scientific associations for plant protection, that can provide a
forum for studying plant health problems in the countries, the subregions and the
i. The participation of farmers' organizations, field workers and the rural population
in campaigns to control pests and diseases, as well as in quarantine measures.
The Heads of Plant Protection of IICA Member States in the Caribbean met in San
Jose, Costa Rica from 15-17th August, 1979, and again from July 27-29th, 1980 in
Barbados. The objectives of these Meetings were to formulate a plan of action for the
Caribbean within the Hemispheric Plant Protection Programme.
The Meeting in Barbados had as its objectives:
1. To analyse the programme objective to make them more precise, more limited in
scope and more realistic in relation to the financial resources of IICA.
2. To establish lines of priority from among the various proposals made at the Meet-
ing in Costa Rica.
3. To establish mechanisms for coordination with Regional and International Plant
The result of this Meeting formed the basis for the orientation of the Programme at
the Regional level. The priorities identified included:
1. Training courses in Plant Quarantine and General Plant Protection.
2. Strengthening post entry Quarantine facilities.
3. Control and eradication of new pests and diseases.
4. Establishment of a Society for Plant Protection in the Caribbean.
5. Establishment of a Regional Newsletter.
The programme recognized the existence of several institutions concerned with plant
protection in the Caribbean. These include:
-The Commonwealth Institute of Biological Control with its track record in the
biological control of pests.
-The Caribbean Agricultural Research and Development Institute (CARDI) with
its work in research and its outreach activities in several of the Islands.
-The Faculty of Agriculture of the University of the West Indies with its research
and teaching capabilities.
Caribbean Conference 9
-Plant Protection divisions of the various Ministries of Agriculture.
The programme, however, recognized that there was no agency that provided a
formal mechanism for coordination and cooperation in plant protection and that recip-
rocal technical cooperation which is so vital in the region because of the lack of plant
protection capability in some of the smaller territories and the limited human and finan-
cial resources available was not being fostered.
The programme also responded to the need for (1) information on pest and disease
control and (2) lack of professional stimulation among professionals in Ministries of
Agriculture, lack of access to Scientific journals and lack of trained sub-professionals in
plant protection and plant quarantine.
In recognition of these challenges, the following are some of the achievements to
TRAINING-The programme recognizes that the improvement of human resources rep-
resent one of the most important mechanism for the enhancement of agricultural de-
velopment. Consequently, training was given high priority in the actions which were
carried out. There were three types of training provided:
a. Plant Quarantine Training
Effective Plant Quarantine is necessary for the safe movement of agriculture pro-
duce in Regional and International trade. The Heads of Plant Protection in the Carib-
bean recognized that there is an urgent need for trained plant quarantine inspectors in
the Region. Consequently, a Regional Plant Quarantine Training Course was estab-
lished. The course was held in Trinidad and Tobago in 1982 and in Barbados in 1983.
The course objectives were as follows:
1. To develop and foster among Plant Quarantine Inspectors of the Region an aware-
ness of their mutual responsibility to keep the Caribbean free from foreign pests and
2. To improve the skills of Plant Quarantine Inspectors in the detection and treat-
ment of plant pests and diseases which pose a threat to Caribbean Agriculture from
either Regional or extra-regional sources.
3. To improve communication between Plant Quarantine Inspectors of various ter-
ritories of the Region.
4. To form the basis for the preparation of a Caribbean Plant Quarantine Training
The course was designed primarily for inexperienced Plant Quarantine Inspectors
and dealt with the general principles of plant quarantine and the duties, responsibilities
and requirements of Plant Quarantine Inspectors.
Twenty-one Plant Quarantine Inspectors have been trained so far. This include six
from Barbados, two from Dominica, two from Grenada, two from Guyana, one from
Haiti, two from Jamaica, one from St. Kitts, one from St. Lucia, two from Suriname
and six from Trinidad and Tobago.
b. Integrated Pest Management
In collaboration with Caribbean Agricultural Research and Development Institute
(CARDI), the University of the West Indies Faculty of Agriculture and the Consortium
for International Crop Protection (CICP) a two week training programme on Integrated
Pest Management was held at the Faculty of Agriculture, University of the West Indies,
St. Augustine, Trinidad and Tobago 10th-21st, 1981. The seminar was attended by
twenty-one participants from thirteen countries in the Region and together with partic-
ipants from Trinidad and Tobago there was an average daily attendance of thirty-five
persons. Lecturers were drawn from the Consortium for International Crop Protection,
the University of the West Indies, Ministry of Agriculture of Trinidad and Tobago, the
Commonwealth Institute of Biological Control, the Caribbean Agricultural Research
and Development Institutes, Caroni Limited and IICA.
10 Florida Entomologist 69(1) March, 1986
The course dealt with the basic concepts of integrated pest management and their
application to the solution of pest and disease problems of crops grown in the Caribbean.
While most participants expressed a high level of satisfaction with the training prog-
ramme, it was the opinion of all that a more sustained effort and long-term training in
this area was necessary. Certificates were presented at the end of the course.
c. Specialized Training
1. Training for the detection of pests in containerized cargo-Two officers of the
Ministry of Agriculture, Lands and Food Production were training in this subject area.
2. Training in Acarology-An officer of the Ministry of Agriculture, Lands and Food
Production was trained in Acarology.
3. Training in postharvest losses-Provided to an officer of a Research Institute in
Trinidad and Tobago and in seed pathology for officers in Jamaica.
INFORMATION-In several of the Islands, access to current scientific literature is lack-
ing; therefore, the programme introduced The Caribbean Plant Protection Newsletter
in 1981. The Newsletter, which is distributed annually, collects relevant information
from the scientific journals and from research in the region and makes it available in
the form of abstracts to plant protection personnel throughout the region. It publishes
information on the following:
1. New research findings in the Caribbean.
2. New research findings elsewhere which are relevant to the region.
3. Abstracts of relevant literature.
4. Information on conferences and meetings.
5. Activities of plant protection personnel and institutions in the Caribbean.
6. IICA's Regional Plant Protection Programme.
7. Information on new equipment, pesticides, etc.
8. Feature articles on regional plant protection matters.
A new feature has recently been introduced where readers of the newsletter can
obtain photocopies of the original articles from the editor.
In addition to the newsletter the programme has published a document entitled "A
Bibliography of Plant Disease Investigations in the Caribbean from 1880-1980". This
publication, which contains references to cover 3,000 publications on plant disease work
in the Caribbean, is intended to serve as a reference source for Plant Pathologists,
Nematologists, Agronomists, Research students and other persons interested in Agri-
culture in the Caribbean.
The Bibliography covers all aspects of plant diseases and their control, including
diseases caused by fungi, bacterial, viruses and nematodes. The geographic area co-
vered by the references includes all the territories of the Commonwealth Caribbean.
The material covered by the Bibliography dates from the 1880's to 1980. The entries
were compiled mainly from primary sources, many of which have not been indexed
previously. It would be impossible to list all the titles searched; however, the main
sources of information were Agricultural News, Tropical Agriculture, West Indian Bul-
letin, the Journal of the Jamaica Agricultural Society, the Journal of Agricultural Soci-
ety of Trinidad and Tobago, and the Annual Reports and Publications of the Depart-
ments and Ministries of Agriculture throughout the Region. The Secondary sources
included Regional bibliographies on agriculture and related topics and computerised
literature searches of the Commonwealth Agricultural Bureau data bases. Copies of this
bibliography have been sent to all plant protection personnel in the Caribbean.
A book on Plant Disease Diagnosis was prepared by the Specialist to provide plant
protection personnel with an introductory handbook for the practical study of plant
diseases. The book outlines the basic steps, facilities and procedures which are neces-
sary for the accurate diagnosis of plant diseases. This publication is especially valuable
Caribbean Conference 11
to laboratory assistants who are being introduced to Plant Pathology for the first time,
to give them the rudimentary principles and techniques which are required for plant
THE ESTABLISHMENT OF A MECHANISM FOR PROFESSIONAL STIMULATION-The
establishment of a society for plant protection in the Caribbean in 1981 responded to the
needs of the region for a professional body to give professional stimulation and also to
act as a mechanism for communication and coordination of plant protection activities.
The society which now has some 121 members throughout the region has as its objec-
tives the following:
-to strengthen inter-governmental and inter-institutional cooperation in plant pro-
tection in the Caribbean;
-to establish a forum for the discussion of plant protection issues affecting Carib-
-to act as a forum for the exchange of ideas and information among plant protection
personnel in the Caribbean;
-to promote and stimulate research and teaching in plant protection subjects, viz.,
Entomology, Plant Pathology, Weed Science, etc. and to ensure that these are integ-
rated into the discipline of plant protection;
-to stimulate discussion and actions to ensure that the Caribbean environment
remains free from contamination by pesticides;
-to carry out all other activities which may be associated with preserving the plant
genetic resources of the Caribbean from destruction by pests and diseases as may be
defined by the Executive Committee.
IICA assisted in the establishment of the Society for Plant Protection in the Carib-
bean at the Second Regional Meeting of Plant Protection in Latin America and the
Caribbean held in Mexico City in October 1980.
The Society has held two meetings so far. In 1981, it met in Jamaica and discussed
urgent plant pests and disease problems in the Caribbean. In 1983, it met in Trinidad
and Tobago and discussed the harmonization of pesticides legislation in the Caribbean.
A document describing the Society has been prepared and is available.
TECHNICAL SUPPORT-This involves the direct use of the Specialist's expertise in the
assistance to the Member States. It involves the answers given to questions of a plant
protection nature and it provisions a resource person to assist in the solution of Regional
problems. Examples of this would be the assistance given to the Barbados Ministry of
Agriculture in the preparation of a Plant Quarantine facility for Cocoa; the advice given
to Jamaica on the re-organization of the Plant Quarantine System, and on the design
and analysis of plant protection experiments; suggestions for strengthening Plant
Quarantine in Guyana; diagnosis of diseases of sugarcane in Haiti and; suggestions for
studies on yellow spot disease of sugarcane, Caroni Limited Trinidad and Tobago.
In addition to these specific areas of work a number of studies have been carried
out in collaboration with regional and extra-regional institutions. The following are
examples of studies carried out:
a. An analysis of Plant Quarantine Systems in the Caribbean in collaboration with
Animal and Plant Health Inspection Services (APHIS).
b. Training and Research needs in seed pathology in the Caribbean in collaboration
with the Danish Institute for Seed Pathology in developing countries located in
c. Study of the pest risks associated with the movement of agricultural produce
between St. Vincent, Grenada and Trinidad and Tobago, (in collaboration with Dr. G.
V. Pollard of the University of the West Indies).
d. Study on the economic impact of Moko disease on the economy of Grenada, (in
collaboration with Dr. G. V. Pollard of the University of the West Indies).
12 Florida Entomologist 69(1) March, 1986
These studies were designed to generate new information necessary for the im-
plementation of plans within the programme.
RECIPROCAL TECHNICAL COOPERATION-RTC means using the technical skills avail-
able in some countries to benefit others through IICA's action as a means of reciprocal
transfer of know-how, and the exchange of technical personnel and useful experience.
The International Agency acts only to facilitate and finance in the relationship. This
was used successfully during the programme in the following activities:
1. Assistance to Grenada in the Control of Thrips of Cocoa-Thrips have been recog-
nized as a severe problem in cocoa production in Grenada. The Government sought the
assistance of IICA and Dr. Eslie Alleyne, Entomologist, Ministry of Agriculture, Bar-
bados was contracted by IICA to provide technical assistance in Grenada. Dr. Alleyne's
recommendations for the control of thrips in Grenada have been taken seriously by the
Ministry of Agriculture authorities and the Chief Plant Protection Officer has now been
seconded to Grenada Cocoa Association to manage the thrips problem on a full-time
2. Assistance to the Food and Agriculture Corporation of Trinidad and Tobago in
the Assessment of Pest Problems of Banana at Orange Grove-The Food and Agricul-
ture Corporation of Trinidad and Tobago, an Agency set up for corporate action in the
agricultural sector under the aegis of the Ministry of Finance and Planning is cooperat-
ing with the Orange Grove National Sugar Company of Trinidad and Tobago in the
development of a banana enterprise geared to the production of green bananas either
for fruit market or use as a basic carbohydrate food item to partially displace rice and
other wheat-derived staples. Based on discussions with the Inter-American Institute
for Cooperation on Agriculture (IICA) for close cooperation on technical matters, the
Corporation sought the assistance of IICA through its national office in Trinidad and
Tobago to evaluate the incidence of pest and diseases in the existing plantings of banana
at the estate and to develop a strategy for management of these pests and diseases in
the future. The study was carried out during the month of May, 1983. Mr. Frank
McDonald, Ministry of Agriculture, Guyana, was contracted to look at the Moko disease
aspects of the problem.
HARMONIZATION OF PESTICIDES LEGISLATION IN THE REGION-There has been a
dramatic increase in pesticide usage in the Caribbean during the last decade and like
many parts of the world there is concern with respect to the potential human and
environmental hazards of pesticide use. The concept of the harmonization of pesticide
legislation and registration requirements in the region stemmed from the initiatives
developed by the FAO in 1977 to seek to harmonize the legislative aspects of pesticide
use throughout the world. A meeting to consider the harmonization of pesticide legisla-
tion and the registration process in the region was held in Trinidad and Tobago under
the auspices of IICA and with financial assistance from the National Agricultural Chem-
ical Association (NACA) and GIFAP.
The meeting considered the status of pesticide legislation in the region and estab-
lished guidelines for harmonization. The meeting had as its objectives:
-to examine and analyse the status of pesticide legislation in the Caribbean;
-to become familiar with measures currently in operation and to recommend others
for standardizing the diverse requirements for pesticide use;
-to provide guidelines to those countries that need to enact legislation to regulate
the use of guidelines, norms and technical procedures (chemical, analytical, biological,
-to examine the status of establishment or acceptance of the maximum residue
levels of pesticide (tolerance) in food in each country;
-to analyse the environmental impact of the large scale application of pesticides in
Caribbean Conference 13
-to stimulate training in pesticide registration procedures and in the safe and effi-
cient use of pesticides.
Forty-five delegates from throughout the Caribbean and from Regional and Interna-
tional Organizations attended the meeting.
The meeting recommended, among other things, that the Pesticide and Toxic Chem-
icals Act of Trinidad ahd Tobago should be used as a model in the formulation of regional
pesticide legislative actions. Follow-up action to this is now being taken in the develop-
ment of a Regional Training Programme in pesticide safety in collaboration with the
World Bank and the University of Miami, School of Medicine. A 263 page document has
been prepared as a result of the meeting.
A MECHANISM FOR COORDINATION AND COOPERATION-THE HEADS OF PLANT
PROTECTION MEETING-The Heads of Plant Protection Meeting which has been held
annually since 1981 provided an excellent mechanism for consultation, cooperation and
communication in plant protection in the region.
The meetings which are held in a different country each year provide opportunities
for the Heads of Plant Protection to visit the countries of the region and to observe the
programmes and facilities which exists in the region. The meetings contribute to:
1. Reduction in the possibilities for duplication of efforts.
2. Establishment of bilateral cooperative programme in the region.
3. Establishment of a fraternity of plant protection which can be a very important
asset in problem solving.
4. It give guidance to the region programme.
5. Provides a forum for coordination of all technical assistance programmes in the
DEEPENING OF THE SCOPE OF THE PROGRAMME BY DIRECT ASSISTANCE TO NA-
TIONAL SYSTEMS-The scope of the programme has now been deepened by the estab-
lishment of national professionals in plant protection in the IICA offices in Dominica,
St. Lucia and Grenada. In addition, in collaboration with USAID and USDA/APHIS
there are proposals to set up a pest management unit in Grenada. This initiative re-
sponds to the need for an effective plant protection capability in the small island states
of the region which are embarking on a major thrust in the development of fruits and
vegetables for the export markets of North America and Europe. The initiative also
responds to the following situation and I quote from a recent document "Although
chemical pesticides-particularly insecticides and herbicides-are being utilized in in-
creasing levels, the pest problems actually seem to be worsening. Annual pest losses
in cocoa now approach an estimated $1 million (U.S.). Leaf spot disease, Moko disease,
and root knot nematodes cause crippling losses in banana if expensive chemical treat-
ments are not applied. GCA and GBCS together annually spend $450,000-about 25
percent of their total budget-for control of cocoa and banana pest". It also responds
to need to have in place a point of contact for the delivery of technical cooperation in
plant protection from International agencies.
In Suriname, a specialist in coconut pests and diseases has been located in the IICA
Office in that country to attend to the pressing problems of coconut mainly "Hart Rot
disease" and Castnia spp. It is hoped that with assistance from other agencies and
friendly governments that these iniatives will result in the development of a regional
centre for investigations on coconut and oil-palms.
1. Preparation of project proposals for a survey of fruitflies in the Caribbean.
2. Preparation of field guides to plant pests and diseases of importance in the Caribbean
and bibliography of pests.
14 Florida Entomologist 69(1) March, 1986
3. Development of Third Regional Plant Quarantine Training Course.
4. Survey of the incidence of mango seed weevil in the Caribbean.
5. Workshop on the detection of pests and diseases of fruits in the Caribbean.
6. Establishment of Regional Training Programme in Pesticide Safety.
7. Establishment of a Data Base in Plant Protection for the Caribbean.
ALLEYNE, E. A Report on a visit to Grenada to advise on the control of cocoa thrips
(Solenothrips rubrocinitus) on cocoa. 1981. Mimeo 16 pp.
ANON. An Investigation of the Incidence of Pest and Disease at Orange Grove National
Sugar Company, Port-of-Spain, Trinidad and Tobago, IICA Office, 1983. 13 p.
ANON. The Society for Plant Protection in the Caribbean. Its origin, Constitution and
Current Membership. Port-of-Spain, IICA Office, 1982. 19 pp.
BRATHWAITE, C. W. D. An Introduction to the Diagnosis of Plant Disease. Inter-
American Institute for Cooperation on Agriculture. Series Book and Education
Materials No. 47, 1981. 49 p.
BRATHWAITE, C. W. D. Perspectives for Plant Protection in the Caribbean. Port-of-
Spain, Trinidad & Tobago, IICA Office, 1981.
BRATHWAITE, C. W. D., ALCOCK, M. AND SOODEEN, R. A bibliography of Plant
Disease Investigations in the Commonwealth 1880-1980. Inter-American Insti-
tute for Cooperation on Agriculture Miscellaneous Publication No. 328. 1981. 280
BRATHWAITE, C. W. D. and POLLARD, G. V. The essential role of pest and disease
control in crop production in the Caribbean. Agricultural Extension Newsletter
(Trinidad & Tobago). 12: 32. 1981.
BRATHWAITE, C. W. D. The challenge for Plant Protection in the Caribbean in the
1980's and beyond. In Meeting of the Society for Plant Protection in the Carib-
bean. 1st, Kingston, Jamaica, 1981. Urgent Plant Pest and Disease problems in
the Caribbean. Edited by Chelston Brathwaite and Gene Pollard. Inter-American
Institute for Cooperation on Agriculture Miscellaneous Publication No. 378. 1982.
BRATHWAITE, C. W. D. Crop Protection in the 1980's-an analysis of present alterna-
tive technologies. In New Technologies in Food Production. Port-of-Spain,
(Trinidad & Tobago). 12: 32-36. 1981.
BRATHWAITE, C. W. D. IICA'S activities in the Caribbean. Port-of-Spain, (Trinidad
and Tobago), IICA Office, 1983 6 p.
BRATHWAITE, C. W. D. Pest and Diseases of Onion. In Workshop on Onion Production
and Research for the Eighties, Bridgetown, Barbados, 1983. Proceedings. Inter-
American Institute for Cooperation on Agriculture Miscellaneous Publication No.
378. 1982. 260 p.
CARIBBEAN PLANT PROTECTION NEWSLETTER. Vol. 1 Nos. 1-4, 1981-1984. Edited
by Chelston W. D. Brathwaite. Port-of-Spain, (Trinidad and Tobago), IICA Of-
Meeting of the Society for Plant Protection in the Caribbean, 1st Kingston, Jamaica,
1981. Urgent Plant Pest and Disease problems in the Caribbean. Edited by
Chelston W. D. Brathwaite and Gene V. Pollard. Inter-American Institute for
Cooperation on Agriculture Miscellaneous Publication No. 328. 1982. 260 p.
Meeting on the Harmonization of Pesticide Legislation in the Caribbean, Port-of-Spain,
(Trinidad and Tobago), 1983. Proceedings. Edited by Chelston W. D. Brathwaite.
Inter-American Institute for Cooperation on Agriculture Miscellanous Publica-
tion No. 379. 1984. 253 p.
POLLARD, G. V. The Economic Impact of Moko Disease on the Economy of Grenada.
Port-of-Spain, (Trinidad and Tobago), IICA Office, 1983. 14 p.
POLLARD, G. V. 1983. Report on a Visit to Grenada, St. Vincent and St. Lucia to
investigate the Potential Pest Risks Associated with the Movement of Agricul-
Caribbean Conference 15
tural Produce via the Inter-island schooner trade. Report to IICA, Port-of-Spain,
Trinidad, November 1983. 23 pp.
SMALL, L. W. The legal framework of plant quarantine systems in the Caribbean.
AN ENVIRONMENTAL TOXICOLOGY PROGRAM
D. L. SHANKLAND
Department of Entomology & Nematology
University of Florida
Gainesville, Florida 32611
H. N. NIGG
Citrus Research and Education Center
700 Experiment Station Road
Lake Alfred, Florida 33850
The total burden of hazardous waste and pesticides in Florida does not appear exces-
sive when compared with national figures. However, Florida is particularly vulnerable
to these materials because it relies heavily on ground water for drinking water.
Florida has enacted an elaborate set of laws on water quality, hazardous waste
management and disposal, pesticide usage, fuel storage, landfill management, and re-
lated matters that bear upon water and general environmental quality. In many cases,
however, there are no cost effective technologies available that have been approved by
the responsible agencies to permit compliance with these laws. This obvious need for
strong research and delivery programs is being met by the Institute of Food and Agri-
cultural Sciences at the University of Florida, the Center for Biomedical and Toxicolog-
ical Research at Florida State University, and the Florida Toxicological Research
Center at the University of South Florida. These agencies coordinate their efforts
through the Center for Environmental Toxicology. The Toxocological Research Coor-
dinating Committee ensures maximal contribution of these programs to the state
through annual assessment and reports to the Governor and the Legislature.
A basis for research and development, thus, has been established in law and in
program development. Fruitful results and real benefit to the state will come only with
realistic support. To date funding has been inadequate for supporting the expanded
research mandated by law.
La carga total de desperdicios peligrosos y de pesticides en la Florida no parece ser
excesiva cuando se comparan con los datos nacionales. Sin embargo, la Florida es par-
ticularmente vulnerable a estos materials porque depend much del agua debajo de
la superficie de la tierra para beber.
La Florida ha enactado una series de elaboradas leyes sobre la calidad del agua, la
administraaci6n y disposici6n de desperdicios peligrosos, uso de pesticides, al-
macenamiento de combustible, administraci6n de rellenos de tierra, y materials re-
lacionadas que tienen que ver con la calidad general del agua y del medio ambiente. Sin
embargo, en muchos casos no hay tecnologia disponible que su costo sea efectivo y que
haya sido aprobado por agencies responsables que permit obedecer esas leyes. La
16 Florida Entomologist 69(1) March, 1986
obvia necesidad de programs de investigaci6n y de rendici6n, esta siendo llenada por
el Institute de Alimentos y Ciencias Agricolas de la Universidad de la Florida, el Centro
de Investigaci6n Biomedico y Toxicol6gico de la Universidad del Estado de la Florida,
y el Centro de Investigaci6n Toxicol6gico de la Florida en la Universidad del Sur de la
Florida. Estas agencies coordinan sus esfuerzos a traves del Centro de Toxicologia
Ambiental. El Comit6 Coordinador de Investigaciones Toxicol6gicas aseguran la max-
ima. contribuci6n de estos programs al estado a trav6s de evaluaciones anuales y
reports al Gobernador y a la Legislatura.
De aqui que se ha establecido en ley y en desarrollo de program una base de
investigaci6n y desarrollo. Resultados fructiferos y beneficios reales para el estado
vendran solo con apoyo realistic. Hasta ahora, apoyo monetario ha sido inadecuado
para mantener y expandir las investigaciones mandadas por la ley.
Under the Resource Conservation Recovery Act of 1976 (RCRA-Public Law 94-580),
the United States Environmental Protection Agency (EPA) is required to institute a
national program to control hazardous wastes. It was the intent of Congress in passing
RCRA that states assume responsibility for controlling hazardous waste within their
borders. In 1980 the Florida legislature passed a comprehensive Florida hazardous
waste management act which codified into state statutes the federal requirements estab-
lished by EPA and RCRA.
There are in the world approximately five million known chemical compounds of
which about 70,000 are in commercial use (Fishbein 1980). EPA has named 654 of these
chemicals as hazardous materials that pose special problems if disposed of into the
environment (CFR 40, Chapter 1, Part 261.33). They are classified as hazardous because
of toxicity, reactivity, ignitability, or corrosivity. Two hundred and sixty of these com-
pounds are listed for their acute toxicity and many of the pesticides in the list are among
this latter group. According to the law, these materials become hazardous wastes when
they are intended for disposal or acutally disposed of.
EPA estimates that approximately 581 billion pounds of hazardous waste are gener-
ated in the U.S. annually (Ney 1984). On the average, this amounts to almost 158
thousand pounds for every square mile of land and water in the U.S. Furthermore, it
is estimated that about one billion pounds of pesticides are used annually in the U.S.
(Pimentel 1979). These pesticides, on the average, enter the U.S. environment at the
rate of about 270 pounds per square mile, and account for about 0.17% of the total
annual burden of hazardous materials on the environment.
This burden is, of course, not uniformly distributed. Florida is a smaller than average
generator of hazardous waste, but larger than average user of pesticides. In a 1977
study of hazardous waste generators in Florida (Carter 1977), 320 organizations, includ-
ing industrial firms and universities, were surveyed; and it was estimated that they
generated approximately 638 thousand tons of waste per year or about 22 thousand
pounds for every square mile of land and water in the state. Accurate figures on pes-
ticide use in Florida are not available, but the state is generally considered to be second
only to California in that regard. Thus, Florida is burdened with only about one-seventh
the national average of hazardous waste per square mile but more than the national
average of pesticides.
The total burden of hazardous waste and pesticides in Florida does not appear exces-
sive when compared with national figures. However, Florida is particularly prone to
serious problems from these materials because of the nature of its water supply (FDACS
1984). Over 90% of Florida's population relies on ground water for drinking water.
Ground water is easily recharged by rain and surface runoff in much of the state and
is easily contaminated by chemicals from spills, surface discharges, dumps, landfills,
Caribbean Conference 17
and pesticide applications. Florida also shares the national concerns over atmospheric
pollution, acid deposition, mycotoxins in food, and other kinds of environmental pollu-
tion. However, during the past three years the discovery of ground water and well field
contamination by pesticides and other chemicals, the legislative action on hazardous
waste management, and enforcement programs by the Florida Department of Environ-
mental Regulation make it clear that the most urgent concern in Florida is point and
nonpoint source pollution of ground water by toxic organic chemicals. The importance
of this issue was emphasized in the summary of the Report of the Speaker's Task Force
on Water Issues, Florida House of Representatives. This March, 1983, report cited
contamination of ground and surface waters with hazardous wastes, sewage, industrial
wastes, pesticides, and other chemical products as the most serious threat to Florida's
supplies of clean water and its fragile ecosystem. The report further cited the absence
in Florida of systematic programs to either assess the extent of, or minimize or remove
the threats posed by these different classes of pollutants.
These circumstances led to an elaborate set of laws on water quality, hazardous
waste management and disposal, pesticide usage, fuel storage, land fill management,
and numerous other matters that bear upon contamination of water and other parts of
the environment by chemicals. Enforcement agencies, like the Florida Department of
Environmental Regulation and U.S. EPA, are mandated to enforce these laws. In many
cases, however, there are no cost effective technologies available that have been ap-
proved by the agencies to permit compliance with the laws. The only approved method
of disposal of some hazardous waste, including for example large areas of pesticide
contaminated land, is to put the material in sealed drums and ship it to an approved
dump in Alabama. This is not only prohibitively expensive, but it cannot be a permanent
solution. Alabama cannot serve as an infinite sink for wastes from the southeastern
An obvious need for strong research and delivery programs to address these matters
was recognized by the Institute of Food and Agricultural Sciences (IFAS) at the Uni-
versity of Florida, Florida State University, and the University of South Florida some
years ago, before the intrusion of aldicarb and ethylene dibromide into ground water
became public issues, and before other cases of ground water contamination received
public notoriety. We need to develop environmentally sound, cost effective technologies
to dispose of hazardous chemicals, and to decontaminate soil and water that have already
been contaminated by them. We need sound data bases for determining human health
risk posed by chemical pollutants in our water. We need to understand how chemicals
behave in the environment, and we need to know the toxicological implications of these
events. We must have the ability to identify and determine the concentrations of these
chemicals in soil, water, food, and other parts of the environment. Finally, the state
needs experts in epidemiology to assess the effects of environmental pollutants on public
health. Toward meeting these needs, the University of Florida, Florida State Univer-
sity, and the University of South Florida independently organized centers of environ-
The University of Florida researches analytical methods for pesticides, fate and
transport of chemicals in the environment, human exposure to pesticides, water treat-
ment, and many other relevant environmental subjects. These were not, however,
coordinated and funded to address the urgent environmental toxicology needs of the
state. In an effort to provide coordination and funding mechanisms, the Center for
Environmental Toxicology was formed in 1982. The center has missions in research,
extension, and instruction. At the present, the research mission deals mainly with the
development of analytical methods for chemical pollutants, the fate and transport of
those chemicals in the environment, and their toxicology. The extension component
serves as an authoritative source of information on the issues for the public, state
18 Florida Entomologist 69(1) March, 1986
agencies, legislators, or other interested parties. The mission in instruction is primarily
graduate education and training to prepare scientists to work in this important aspect
of environmental science.
At Florida State University, the Center for Biomedical and Toxicological Research
forms the administrative hub for interdisciplinary research on ecological effects of
marine pollutants, effects of toxic organic substances in the environment, effects of
heavy metals, pesticides, and other pollutants on aquatic systems, etc. The center has
provided the Florida Departments of Health and Rehabilitative Services and Environ-
mental Regulation with health risk assessments of carcinogenic and toxic chemicals in
ground water. That center also has a mission in graduate education.
At the University of South Florida there is a large and growing Medical Center, a
newly established College of Public Health, and a close working relation with the VA
Hospital all of which form a strong base for their Florida Toxicological Research Center.
The missions of that center are basic research into the toxic chemicals present in Florida
and investigation into alternative chemicals and processes to presently used hazardous
chemicals, and to provide technical support for Florida state agencies with analytical
expertise and scientific consultation.
In recognition of the serious nature of the threat by chemical pollutants to Florida's
water supply and to the welfare of the citizens of the state, the legislature in 1984
provided a statutory basis for a systematic state-wide research program on those mat-
The Law of Florida, Chapter 84-338, Section 72 charges the Center for Biomedical
and Toxicological Research at Florida State University, the Institute of Food and Ag-
ricultural Sciences (IFAS) at the University of Florida, and the Florida Toxicological
Research Center at the University of South Florida to increase their research on chem-
icals that may adversely affect human health and the environment, and to do so without
unnecessary duplication of effort.
To ensure maximal contribution of these programs to the state, Section 73 of the
law established the Toxicological Research Coordinating Committee (TRCC) made up
of representatives of the three universities, and representatives from other appropriate
universities as recommended by the Board of Regents. The Committee is chaired by
the representative of the Center for Environmental Toxicology. The Committee is
charged to ensure efficient use of the state's resources, and toward that end, to meet
at least once a year to review research, develop activities, and establish priorities as
determined by state needs. The Committee is to submit annual reports concerning the
activities of each participating university and short- and long-range plans of each for
protecting human health and Florida's environment. The report shall be submitted on
March 15 to the Governor, the President of the Senate, and the Speaker of the House
Section 74 of the law calls for submission of reports from participating universities
to the Toxicological Research Coordinating Committee on:
1) Chemicals that may affect human health and welfare, including epidemiological
2) Analytical methods, environmental fate and transport, and toxicology.
3) Environmentally safe methods to control pests other than through the use of
The Committee in turn is to provide risk assessment analysis to the Department of
Agriculture and Consumer Services, the Department of Health and Rehabilitative Ser-
vices, the Department of Environmental Regulation, and Pesticide Review Council. It
must recommend standards of safety for chemicals in the environment, and perform
other functions necessary to carry out the provisions of the law. Section 75 of the law
mandates the creation of a data bank on environmental toxicology research results in
Caribbean Conference 19
the Center for Environmental Toxicology in IFAS. Each participating university is to
provide results of completed research on environmental fate and transport and toxicol-
ogy to the data bank. Although not specified in Section 75, the provisions of Section 74
imply that reports should also include results on environmentally safe methods to control
pests other than through the use of chemicals. On September 30 of each year, IFAS is
to publish a listing of publically available studies in the data bank. The list must also
be issued to the Department of Agriculture and Consumer Services, the Department
of Health and Rehabilitative Services, the Department of Environmental Regulation,
and the Pesticide Review Council.
Unfortunately, the legislature was guilty of an oversight. The law was enacted
without funding. Research relevant to the law has been funded by limited amounts of
general review and by extramural funds of various kinds. In all cases, these funds are
inadequate for, or contractually restricted from supporting the expanded research man-
dated by the law.
During the 1985 legislative session the oversight of last year was corrected by the
provision of $750,000 from the Water Quality Trust Fund to support this state-wide
program. That funding is not assured for next year, however. In spite of that, the
Toxicological Research Coordinating Committee is proceeding with plans for the de-
velopment of the Data Bank and with specific research projects selected from a list of
high priority issues developed by the Department of Environmental Regulation, the
Department of Agriculture and Consumer Services, and the Pesticide Review Council.
Requests for support from general revenue for continuation of this support will be
coordinated next year during the legislative session.
It cannot be argued that a ground water contamination crisis exists in the state of
Florida as a whole. The citizens of Fairbanks, Florida, however, whose well field was
made unusable as a source of drinking water by contamination with toxic chemicals from
a mismanaged chemical dump, have already lived through a crisis. That problem is
being solved by extending the Gainesville water system to Fairbanks at a cost in excess
of two million dollars.
The Florida Department of Health and Rehabilitative Services has been analyzing
well water, mainly from drinking water wells, for the presence of ethylene dibromide
(EDB) in areas where that chemical has been used for nematode control. As of June of
this year, 9,208 wells had been analyzed and 1,019 (11%) of them were contaminated
with ethylene dibromide in excess of the 0.02 parts per billion action level established
by EPA. In fact some wells contained 600 to 700 parts per billion of EDB and the
average among contaminated wells was 6.5 parts per billion. The owners of those wells
face reduced property values and the inordinate inconvenience of relying on bottled
water for household use. The state of Florida has assumed responsibility for some of
these contaminated wells, and it was estimated in 1984 (DER, 1984b) that it would cost
the state $4.9 million to meet that responsibility. At the time of that estimate, however,
solutions to the problem were in terms of possibilities only and included carbon filters,
new wells, and connection to community water supplies. It is probable that the ultimate
cost to the state will be much greater than that estimated.
Data is still being collected by state agencies and Union Carbide Corporation on the
extent to which aldicarb (Temik) has intruded into ground water in areas where it has
The Florida Department of Environmental Regulation is monitoring 402 sites at
which hazardous wastes are being generated (DER, 1984a). Ground water contamina-
tion in excess of adopted standards, or in some cases by specific contaminants for which
standards have not been adopted, has been confirmed at 119 (30%) of those sites. Fur-
thermore, according to DER all 654 of the chemicals on the EPA hazardous materials
list are to be found among hazardous wastes produced in Florida.
20 Florida Entomologist 69(1) March, 1986
Within the constraints of presently available technology, the contamination of wells
with ethylene dibromide is for practical purposes irreversible in situ. That is true also
of wells or aquifers that have become contaminated with gasoline, as happened recently
in a small community southwest of Gainesville, or with other toxic organic chemicals.
Except for activated carbon filtration there are no other proven cost effective
technologies for the homeowner or community to purify water that has become contami-
nated with these chemicals.
Much of the ground water contamination that has occurred in Florida has been due
to mismanagement and irresponsibility. However, the unavailability of cost effective
technologies to dispose of hazardous waste means the accumulation of large amounts of
these materials in storage and increases the probability of accidental or purposeful
illegal disposal of them.
In any event, the need for research and development in these and related matters,
as mentioned earlier, is obvious. A basis for that research has been established in law
and in program development at the universities. Fruitful results and real benefit to the
state will come only with realistic support for those programs. That support will be
sought from various sources; but, in the interest of continuity and to assure that urgent
matters in the state will be addressed, a significant portion of it should come from the
state. Whether or not there is serious intent by the legislature in that regard should
be apparent by this time next year.
CARTER, C. E. 1977. Hazardous Waste Survey for the State of Florida. Prepared for
the Florida Department of Environmental Regulation.
DER. 1984a. The Sites List. Summary Status Report, July 1, 1984-December 31,
1984. Florida Department of Environmental Regulation, Bureau of Operations,
DER. 1984b. Florida's Environmental News. Vol. 6, No. 9. Florida Department of
FDACS. 1984. Summary of soil, hydrogeological and other environmental conditions
in the state of Florida related to pesticide use. Report submitted to the U.S.
Environmental Protection Agency by Florida Department of Agriculture and
Consumer Services, Bureau of Product Evaluation, May, 1984.
FISHBEIN, L. 1980. Potential industrial carcinagenic and mutagenic alkylating agents.
Pages 329-363 in D. B. Walters, ed. Safe Handling of Chemical Carcinogens,
Mutagens, Teratogens, and Highly Toxic Substances, Vol I. Ann Arbor Science,
Ann Arbor, Mich.
NEY, R. E., JR. 1984. Report at National Workshop on Pesticide Disposal, Jan. 28-29,
Clarion Hotel, Denver.
PIMENTEL, D. 1979. A cost-benefit analysis of pesticide use in U.S. food production.
Pages 97-149 in T. J. Sheets and D. Pimentel, eds. Pesticides: Contemporary
Roles in Agriculture, Health, and Environment. Humana Press, Clifton, N.J.
COOPERATIVE INDUSTRY EFFORTS WITH DEVELOPING
COUNTRIES TO IMPROVE AGROCHEMICAL REGISTRATION,
LABELING AND EDUCATION AND TRAINING PROGRAMS
WILLIAM L. HOLLIS
Director of International Affairs
National Agricultural Chemicals Association
1155 Fifteenth Street, NW
Washington, D.C. 20005
The agrochemical industry is cooperatively engaged in carrying out education and
training activities to eliminate misuse and achieve safety in the use of their products
throughout the world. Meaningful and lasting success of these activities will come only
after a sound regulatory foundation for the registration, labeling, and use of agrochem-
icals is established by local governments. This goal is being pursued in Latin America
and the Caribbean as a cooperative effort of governments and industry utilizing a con-
The International Group of National Associations of Agrochemical Manufacturers,
in 1979, began an intense process of consultations with some Latin American gov-
ernments. Formal governmental consultations, under the sponsorship of IICA, fol-
lowed in 1982-83 which included all governments of Latin America and the Caribbean.
As a result recommendations were agreed upon amomg governments for registration
requirements, labeling, toxicity categories, and other safety and training considera-
tions. As of 1984, 11 Latin American nations had accepted the recommendations and 7
others were in various stages of the legislative process.
Most Caribbean nations are legally unable to accommodate the recommendations until
they establish the prerequisite legislation. IICA is preparing a working document for
the governments of the Caribbean to use to establish their own regulations.
An informal consultation between industry and environmental groups to resolve
problems associated with the safe use of agrochemicals and the status of the Food and
Agriculture Organization (FAO) Draft Code of Conduct on the Distribution and Use of
Pesticides are also mentioned.
La industrial agroquimica esta cooperativamente envuelta en llevar a cabo ac-
tividades de educaci6n y entrenamiento para eliminar el mal uso y obtener seguridad
en el uso de sus products a trav6s del mundo. El 6xito duradero y que signifique algo
de estas actividades, vendra solo despues de una s6lida fundaci6n de regulaciones para
la registraci6n, marcar, y el uso de agroquimicos sean establecidas por los gobiernos
locales. Esta meta se esta tratando de llevar a cabo en latinoam6rica y en el Caribe
como un esfuerzo de cooperaci6n de gobiernos e industries utilizando un process consul-
En 1979, el Grupo Internacional de las Asociaciones Nacionales de Fabricantes Ag-
roquimicos, comenz6 un intensive process de consultas informales con algunos gobiernos
latinoamericanos. Durante 1982-83, consultas formales con los gobiernos fueron au-
spiciadas por IICA que incluyeron todos los gobiernos de latinoamerica y del Caribe.
Como resultado, se acord6 entire los gobiernos recomendaciones de requisitos de regis-
traci6n, marcas, categories de toxicidad, y otras consideraciones de seguridad y en-
trenamiento. A partir de 1984, 11 naciones han aceptado las recomendaciones y otras 7
estan en varias etapas del process legislative.
La mayoria de las naciones del Caribe no pueden legalmente acomodar las recomen-
daciones hasta que ellas establezcan las pre-requeridas leyes. IICA esta preparando un
Florida Entomologist 69(1)
document con el cual los gobiernos del Caribe puedan usar para trabajar y establecer
sus propias regulaciones.
Se menciona tambi6n una consult informal entire la industrial y grupos interesados
en el medio ambiente para resolver problems asociados con el uso seguro de los ag-
roquimicos y el estado de la Version del C6digo de Conducta Sobre la Distribuci6n y
Uso de Pesticidas de la Food and Agriculture Organization (FAO)
An international movement to harmonize pesticide registration requirements accord-
ing to FAO recommendations (FAO 1982) is being successfully implemented in Latin
America and the Caribbean as a cooperative effort of governments and industry using
the consultative process (GIFAP/WICEM 1984).
As a focal point there are a few premise statements of obvious facts on plant protec-
tion and some lesser known facts about the agro-chemical industry.
(1) Agricultural food and fiber production is the primary industry of man. It serves
his basic health and economic needs.
(2) There are no major agricultural crops grown anywhere in the world that are free
from pest destruction.
(3) Agrochemicals are the most widely used means of providing immediate and
economic plant protection.
(4) All plant protection schemes and materials, natural and synthetic, have inherent
limitations for pest control that ultimately determine the extent and nature of
their practical use.
In essence, because plant protection is essential to assure food and fiber production
worldwide, and since control options are not infinite, any inherent limitations in efficacy,
safety, or efficiency for whatever controls are being used (Hollis 1977) must be overcome
by correct management in use.
Regarding the agrochemical industry, one of its lesser known features is its size. It
is quite small in the United States as well as internationally. In the U.S. it accounts
for approximately two percent of the gross sales of the U.S. chemical industry (Hollis
1983). Its total worldwide value in sales in 1984 was 13.8 billion U.S. dollars. Even with
recent increases, the total sales value of the worldwide agrochemical industry would
still be less than the documented gross sales of some of our U.S. corporations. For
instance, the EXXON Corporation reported gross sales for 1984 of 97.3 billion U.S.
According to Milton Russell (1984), Assistant Administrator for Policy, Planning
and Evaluation, EPA, the number of major U.S. pesticide producers declined from
approximately 80 companies to 30 companies between 1970-81. The number of com-
pounds screened in relation to the number successfully registered went from 6500/10 in
1967-70 to 82600/11 in 1979. The time from discovery to full registration went from 68
months to 94 months for the same period.
Frawley (1961) cited National Academy of Science figures for 1956 as being three
thousand chemicals screened per one marketable product. He approximated the de-
velopmental cost, in the 1950's, at almost two million dollars per product. The approx-
imate developmental cost per product reported by some industry members today is in
the vicinity of forty million dollars and only one of every 20,000 compounds screened
has a possibility of reaching the marketplace. The research and development cost
excludes the millions needed to build a plant to produce the product.
The global industrial resource of innovative research and development in plant pro-
tection chemistry is shrinking and is considered to be limited now to some 30-40 remain-
ing companies worldwide who still have the financial resources and scientific capabilities
Caribbean Conference 23
to continue. The evolution of newer plant protection products and their establishment
in plant protection field practices takes longer and occurs at a reduced frequency from
fewer sources than in the past. If the agrochemical options available for dependable
plant protection slowly diminish in light of the need for enhanced productivity and crop
diversity to meet the demographer's predictions for the world population in the next
20-30 years, then the potential for a future crisis in plant protection becomes substan-
It is prerequisite to their continual availability that measures be taken to control
the limitations of present and future plant protection products. These limitations are
basically safety and misuse. Measures to overcome such limitations are underway within
the National Agricultural Chemicals Association (NACA) and through our international
organization, The International Group of National Associations of Agrochemical Man-
ufacturers (GIFAP). The strategy followed is the orderly procedure depicted in Figure
1; the Circle of Safety.
For cooperation, there is first the question, how does a regulated industry, regulated
more or less in every country, establish a cooperative effort with Government regulators
and with non-governmental organizations (NGO). To do this, there are certain condi-
tions that must evolve; the demonstration by the established industry of its scientific
^ p OF SA^> C44
Fig. 1. Plant Protection Chemicals-Circle of Safety
24 Florida Entomologist 69(1) March, 1986
competence and credibility, and the willingness to take the initiative to meet with
NGOs, international organizations, and governments in uncompromising forums. Hav-
ing met these conditions, the agrochemical industry can now announce some of their
Within the framework of NACA, the Association has joined in a forum, the Agricul-
tural Chemicals Dialogue Group (ACDG) with a consortium of environmental and church
organizations moderated by the Conservation Foundation. In the past few years, the
ACDG has mutually agreed upon and issued voluntary guidelines for industry on adver-
tising (ACDG 1983) and labeling (ACDG 1985) in developing countries.
The NACA International Registration Committee has given industry support
through the U.S. Department of State to the FAO Draft Code of Conduct on the
Distribution and Use of Pesticides (FAO 1985). The Code had its genesis as a recommen-
dation in the Report of the FAO Consultation on the International Harmonization of
Pesticide Registration Requirements held in Rome, Italy (FAO 1982). NACA was pres-
ent as a member of the GIFAP delegation to the Consultation which supported the
Report. GIFAP has since issued position papers indicating support and cooperation on
principles reflected in the Code pertaining to hazardous substances export (GIFAP
1983b), good marketing practices in pesticide export (GIFAP 1985a), and options for
ensuring quality in stored pesticide products (GIFAP 1985b).
The basic attributes of the Code are that it is voluntary; it is to be observed in
countries that are without national laws in regulating pesticide safety or without regis-
tration controls prior to marketing; and it involves consideration for shared responsibil-
ity for safety measures among government officials, industry, importers and dis-
tributors, and users. The Code consists of twelve articles that comprehensively cover
the essential regulatory requirements and safety measures set forth in the Report of
the October 1982 FAO Consultation (FAO 1982).
The Draft Code was approved by the FAO Committee on Agriculture, comprised of
some 94 delegations, in March 1985 and by the FAO Council in June 1985. The Code
will be considered by the FAO Commission in November 1985 at which time it will be
formally adopted (GIFAP 1985c).
There may be some jurisdiction conflicts and difficulties depending on how some
governments conduct their business in agrochemicals. Nevertheless, the potential suc-
cess of the Code depends most importantly on participating countries and local impor-
ters and distributors remaining attentive along with the industry, to carrying out their
respective responsibilities as identified in the Code (GIFAP 1985c).
The agrochemical industry demands reasonable and responsible regulation. Industry
prefers not to see the Code become a substitute for duly established regulations, but
rather that it serve as an interim measure for safety pursuant to the institution of
appropriate legislation and regulations in those countries to which the Code is relevant.
The GIFAP Latin American Working Group is working cooperatively with the Inter-
American Institute for Cooperation in Agriculture (IICA) to bring this about in the
Industry's cooperative efforts with international bodies and the governments they
represent is accomplished through participation in GIFAP. The agrochemical industry
is one of the few industries that has an international association. It is needed because
national associations have no standing outside their borders. GIFAP is recognized as
the worldwide representative of the agrochemical industry and is accepted by interna-
tional bodies such as United Nations organizations and IICA. GIFAP has for some time
had official status with FAO. Earlier this year, the Director General of the WHO
confirmed the approval of the establishment of official relations between the WHO and
GIFAP (GIFAP 1985d). Such industry recognition must be earned over time by the
demonstration of competence and credibility.
Fiure 2 : World pesticide market
(US $13,000 million end-user sales value in 1981)
excluding non-crop outlets
NORTH AMERICA WESTERN EUROPE OTHERS BRAZIL
AUSTRALASIA INDONESIA INDL
SOVIET UNION AND JAPAN KOREA MEXIC(
WORLD DEVELOPING COUNTRIES
Fig. 2. World Pesticide Market
GIFAP's membership includes over 30 national agrochemical associations which to-
gether comprise more than 950 companies. This membership represents, at the interna-
tional level, more than 90 percent of the world production of agrochemicals. The world
distribution of agrochemicals according to end-user sales value in 1981 is shown in Figure
2. This data was presented (GIFAP 1983b), at a UN interagency meeting in the spirit
of cooperation, to describe the economic and the health and environmental safety consid-
erations that governments of developing countries should consider before deciding to
set up plants to formulate or manufacture agrochemicals.
The member associations of GIFAP have common objectives-to promote crop pro-
tection by appropriate use of agrochemicals worldwide and to ensure that the properties
and application of these products are in conformity with the needs of agriculture and
society; i.e., optimal food and fiber production with minimal hazards for man, animal
To achieve this, some of GIFAP's aims are:
To promote the safe and sensible manufacture, handling, packing and transport of
agrochemicals by setting, and recommending high standards in conformity with interna-
tionally acceptable rules.
To promote the safe and sensible application of agrochemicals, in conformity with
national and international standards and regulations for the protection of the user, the
environment, and the consumer.
To promote harmonization of national and international legislation and regulations
concerning control, testing and approval of agrochemicals.
These aims are in consort with the all-inclusive registration and safety guides in-
itiated at the first such FAO Consultation in 1977 and reported final at the 1982 Consul-
tation (FAO 1982). An examination of the Report (FAO 1982) shows, without question,
that there are benefits for governments, farmers, consumers, and industry by the in-
stitution of the FAO proposals. Meanwhile, the benefits of the FAO proposals remain
Florida Entomologist 69(1)
latent in the absence of government initiative to implement them. GIFAP recognized
this and further recognized that industry is the common denominator in this equation
and the party best able to provide the initiative.
Consequently, GIFAP entered into a unique experiment in 1979 whereby discussions
on regulatory matters with some governments of Latin America, as a group, were made
possible through an informal consultative forum managed and directed by a neutral
non-profit organization, the Policy Sciences Center, Inc. (PSC) interested in public
policy issues. Public funds from the Charles F. Kettering Foundation, the Rockefeller
Brothers Fund, The United Nations Environment Program, and the U.S. Agency for
International Development supported the project.
The Consultative Process (GIFAP 1984a), is a strategy for bringing together parties
of different persuasions, voluntarily, in a neutral forum so as to: a) encourage communi-
cation; b) achieve a better level of mutual understanding of a problems) than existed
before; and c) reach nonbinding consensus agreements that may lead to the resolution
of the problemss. As such, the Consultative Process is a means of legitimizing essential
communications between the regulated industry with its scientific and technological
expertise and governments who need information and cooperation.
The PSC project included an on-site evaluation of the status of considerations given
to labeling, application, and formulation by governments-in eight Latin American coun-
tries. The Review Team, lead by Professor Harvey Cromroy, University of Florida,
issued a Report (Cromroy et. al. 1981) that was reviewed by all participants. Recom-
mendations were presented and when assembled by PSC (GIFAP 1984a), served as the
substance for a final meeting of the Consulation in Key Biscayne, Florida, in 1981. This
experiment in the Consultative Process resulted in:
- candid and respectful dialogue on major concerns to both parties;
- provisions for safety information;
- emphasis on product use and application training;
- clear identification, definition, and organization of main issues and problems; and,
- impetus to undertake formal government consultations.
The industry, in complying with a voluntary commitment to the Forum to provide
safety information, published guides for the safe handling of pesticides in formulating,
etc. (GIFAP 1982), and guides for safe use of pesticides (GIFAP 1983a). Guides pertain-
ing to first aid (GIFAP 1984b) were recently published. All these have been given wide
distribution throughout the hemisphere.
The Consultative Process next shifted from unofficial to official government status
when the Mexican Government, in applauding the efforts of the Key Biscayne Consul-
tation, took the initiative to hold a "Consultation on the Proper Use of Pesticides in
America and the Caribbean" in Mexico City, 1982. The Mexico City Hemispheric Con-
sultation issued proceedings (Direcion 1982) and an official report (GIFAP 1984a) attest-
ing to the essentiality of agrochemicals in food production and recognized problems that
need attention. It identified the benefits that would accrue from the harmonization of
requirements for registration, labeling, and use. The Report (GIFAP 1984a) recom-
mended that IICA conduct Consultations to harmonize pesticide registration require-
ments, recognizing the guidelines set forth by the FAO (FAO 1982), in each of the four
IICA regions; i.e., the Andean Region; the Central Region including Mexico, Panama,
and the Dominican Republic; the Caribbean Region; and the Southern Cone, thus en-
compassing essentially all the nations of Latin America and the Caribbean. Consulta-
tions were held in Cartagena, Colombia, August 1982; San Jose, Costa Rica, April 1983;
Port-of-Spain, Trinidad, August 1983; and Santiago, Chile, August 1983. Reports (IICA
1982, 1983a, b, c) were respectively issued and each included recommendations for
registration requirements, labeling, toxicity categories, and other safety and training
considerations that are universally needed. These were officially approved and signed
Caribbean Conference 27
by all the government delegates.
An open session was held at the beginning of each Consultation to permit a small
GIFAP delegation of experts to present papers (IICA 1982, 1983a, b, c) on registration,
labeling, and toxicology requirements. The registration requirements presented in-
cluded, among other items, suggestions to include information on worker reentry inter-
vals, pre-harvest intervals, and container disposal.
Suggestions for a uniform label format were presented along with the safety idea of
adding a precautionary color band commensurate with the toxicity category of a prod-
uct. The colors, in decreasing order of hazard are: red, yellow, blue, and green. The
colors would be standard according to an international color code and would appear at
the bottom of the label as a band fifteen percent the height of the label. Toxicology
requirements were presented including the suggestion that the WHO classification for
toxicity be used. The suggestions of the GIFAP expert delegation were given favorable
Following completion of all four Consultations, the Director General of IICA pre-
pared a summary report (GIFAP 1984a) of these events for the Inter-American Board
of Agriculture. A Resolution (GIFAP 1984a) to accept the regional recommendations
and implement them "quickly" was subsequently adopted by all the Ministers and Sec-
retaries of Agriculture in Latin America, the U.S., Canada, and the Caribbean at the
October 1983 meeting of the Board in Jamaica.
The status of implementation of the recommendations among the nations of the
hemisphere, as of 1984, includes eleven nations in Latin America who have given full
acceptance and are implementing the recommendations. Seven remaining nations in
Latin America are in various stages of moving through the legislative and regulatory
processes to full acceptance and implementation. Most of the Caribbean nations are
legally unable to accommodate the recommendations until they establish the prerequis-
ite laws. IICA is translating the now established regulations for the Central Region to
serve as a working document for the governments of the Caribbean nations to use to
establish their own regulations.
We are closing in on what was thought to be the impossible-the harmonization of
the regulations of agrochemicals throughout the hemisphere. GIFAP and its member
associations are sincerely encouraged by these progressive and enlightened moves by
the governments of this hemisphere toward the orderly regulation of agrochemicals.
The benefits to be derived from having responsible and reasonable national laws and
regulations for the proper control and safe use of agrochemicals accrue to all concerned
parties: governments, farmers, the public, and the established industry. The benefits
label information and consistent format to meet local needs for correct use and
human and environmental safety;
quality assurance of the product for safety and efficacy reasons;
proper toxicological considerations for safety of all concerned;
user education and training for personal and environmental safety reasons; and
incentives to industry to continue to improve services to local agriculture.
It is not too obvious, but the issue of misuse is addressed throughout these proceed-
ings. Misuse and its attending human and environmental effects does occur and espe-
cially in situations of inadequate registration requirements and procedures as they may
limit regulatory control. Misuse and its effects are always a matter of concern to the
industry and is the main reason we are proponents of sound and enforceable regulations.
The legitimate availability of agrochemicals is a main consideration of the FAO Code of
Conduct (FAO 1985) and of U.S. requirements as evaluated in Congressional testimony
by NACA (NACA 1983).
Meanwhile, developing countries who must import plant protection chemicals and
Florida Entomologist 69(1)
who do not have a regulatory system and basic registration requirements in place are
vulnerable to foreign counterfeiters who fabricate look-a-like agrochemicals which, with-
out quality control, may be contaminated with unknown toxic by-products and perhaps
labeled without precautions. There may be no evidence of efficacy. These products may
be marketed at attractive cut-rate prices directly or through some exporter who may
or may not be interested in following recognized channels of commerce. It is possible
that labels as well as containers may be counterfeits of those of an established company.
That these activities do occur is verified (Deuse 1984). Developing countries must have
the means to evaluate the source and quality of the agrochemicals they import as one
primary means of overcoming inadvertent misuse.
The agrochemical industry continues to be cooperatively engaged in carrying out
education and training activities. It believes that, while these activities are the
mainstays of eliminating misuse and achieving safety, meaningful and lasting success
of these activities will come only after a sound regulatory foundation for the registration,
labeling, and use of agrochemicals is established by local governments (Figure 1).
ACDG. 1983. Guidelines for Advertising Practices in the Promotion of Pesticide Prod-
ucts in Developing Areas of the World. Conservation Foundation, 1717 Mass.
Ave., N.W. Wash., D.C. 20006.
ACDG. 1985. Guidelines on Labeling Practices for Pesticide Products in Developing
Areas of the World. Ibid.
CROMROY, H. L., L. O. ROTH, AND K. J. MOY. 1981. Improving the Safe Use of
Agricultural Chemicals in Latin America. A Research Report on Labeling, Appli-
cation, and Formulation. The Policy Sciences Center, Inc., 270 Broadway Room
1001, New York, N.Y. 10007.
DEUSE, J. P. L. 1984. Imitation and Adulteration of Plant Health Products: A Plague
in Developing Countries. Presented at the Phytopharmacology Seminar, Faculte
des Sciences Agronomiques de l'Etat de Gembloux, Belgium. GIFAP Bulletin
Vol. 11, No. 3, May/June 1985. GIFAP, Avenue Hamoir 12, 1180 Brussels, Bel-
DIRECTION GENERAL de SANIDAD VEGETOL. S.A.R.H. MEXICO CITY. 1982. III
Consultation Meeting on Proper Use of Pesticides in America and the Caribbean.
DIRECTORY. 1982-83. International Group of National Associations of Agrochemical
Manufacturers. P. 11. (Available GIFAP, Avenue Hamoir 12, 1180 Brussels,
FAO. 1982. Report of the Second Government Consultation on International Harmoni-
zation of Pesticide Registration Requirements. AGP: 1982/M/5.
FAO. 1985. International Code of Conduct on the Distribution and Use of Pesticides.
CL87/9-Sup. 1. May 1985.
FRAWLEY, J. P. 1961-62. Process of Discovering and Developing A Marketable Pes-
GIFAP. 1982. Guidelines for the Safe Handling of Pesticides During Their Formula-
tion, Packing, Storage, and Transport. GIFAP, Avenue Hamoir 12, 1180 Brus-
GIFAP. 1983a. Guidelines for the Safe Handling and Effective Use of Pesticides. Ibid.
GIFAP. 1983b. Hazardous Substances Export Policy Position Paper. Ibid.
GIFAP. 1983c. The Manufacture and Formulation of Pesticides in Developing Coun-
tries. Technical Monograph No. 10. Ibid.
GIFAP. 1984a. Agrochemical Industry Briefing Book for the UNEP World Industry
Conference on Environmental Management. Papers with full references on: The
Consultation Process: A Key to Communications, Understanding, Harmonization
Between Governments and Industry; Constraints to Environmental Investment
Caribbean Conference 29
by Agrochemical Industry From Inadequate Propriety Protection; The Role of
Product Stewardship and Co-Shared Responsibility in the Agrochemical Indus-
try. Prepared at the invitation of the WICEM Agenda Committee, 1983. Ibid.
GIFAP. 1984b. Guidelines for Emergency Measures in Cases of Pesticide Poisoning.
GIFAP. 1985a. GIFAP Principles and Objectives of Product Stewardship and Good
Marketing Practices in the Export of Pesticides Position Paper. Ibid.
GIFAP. 1985b. Options for Ensuring Quality in Stored Pesticide Products. Technical
Monograph No. 10. Ibid.
GIFAP. 1985c. International Code of Conduct on the Distribution and Use of Pes-
ticides. Article. GIFAP Bulletin V. 11, N3, May/June 1985. Ibid.
GIFAP. 1985d. Official Relations Between GIFAP and WHO. Article. GIFAP Bulletin
Vol. 11, N2, March/April 1985. Ibid.
HOLLIS, W. L. 1977. The Realism of IPM as a Concept and in Practice With Social
Overtones. Presented at the Annual Meeting of the Ent. Soc. of Am. Symp. IPM
Relative to Fed. Agencies, Academia, and Ind. pp. 12-17. (Available Author,
NACA, 1155 15th Street, N.W. Washington, D.C. 20005).
HOLLIS, W. L. 1983. Agrichemical Residues in Perspective to Agriculture and Food
Risks and Hazards. Assoc. Food & Drug Officials Quarterly Bulletin, Vol. 47,
No. 2, p. 112.
IICA. 1982. Reunion de Consulta Sobre la Armonizacion de Etiquetado Y Registro
de Plaguicidas Para Los Paises Del Area Andina. Proceedings in Spanish. (Avail-
able IICA, Apartado Postal 55 2000 Coronado, San Jose, Costa Rica.)
IICA. 1983a. II Reunion de Consulta Para La Armonizacion de Criterios en Registro
Y Etiquetado de Plaguicidas Para Los Paises Del Area Central. Proceedings in
IICA. 1983b. Pesticide Legislation and the Registration Process in the Caribbean.
IICA. 1983c. Reunion de Consulta Para la Armonizacion de Criterios en Registro Y
Etiquetado de Plaguicidas Para Los Paises Del Area Sud. Proceedings in
NACA. 1983. NACA Position on Pesticide Exports and U.S. Export Regulations.
Testimony before the Subcommittee on Department Operations, Research, and
Foreign Agriculture of the Committee on Agriculture, U.S. House of Rep. June
9. (Available Author, NACA, 1155 15th Street, N.W. Washington, D.C. 20005).
RUSSEL, M. 1984. A Look at EPA's Regulatory Policy. UNEP Industry and Environ-
ment, July/Aug./Sept. 84 ed.
aCtE OF SAFE
- .= I
Florida Entomologist 69(1)
LATIN AMERICAN ENTOMOLOGICAL SERIALS
ANN H. KING
Institute of Food and Agricultural Sciences
University of Florida
Gainesville, FL 32611
This list of nearly 300 serial titles covers the Latin American literature of entomology
comprehensively. Titles in the areas of general zoology plus plant and animal pathology
are also included when they are substantially entomological in content. Serials included
are published by governmental agencies at various levels, as well as commercial firms
and scientific societies.
Latin America has always been an area of great interest to scientists at the Univer-
sity of Florida; consequently, the acquisition of scientific publications from this area has
always had a high priority. This list of entomological serials has been produced as a
means of identifying publications for possible acquisition.
In 1970, a significant list entitled The Serial Literature of Entomology: A Descriptive
Study by Gloria M. Hammack, was published under the sponsorship of the Entomolog-
ical Society of America and the National Science Foundation. It contained a substantial
number of Latin American titles. In 1968, a similar unpublished list was produced
locally by Jane Rayborn of the Florida Department of Agriculture and Consumer Ser-
vices Division of Plant Industry Library (DPI Library). The present list was prompted
by the need to combine the relevant sections of both the Rayborn and Hammack lists
and to incorporate titles begun since 1970.
This list includes titles published by governmental agencies at various levels, as well
as commercial publishing firms and scientific societies. Grouping is by major geographic
area (Carbibbean, Central America, South America) and then by country of origin.
Entry is under the title, except when the publication is produced by a society, govern-
ment agency, or university, in which case the sponsoring agency is generally used. This
type of entry is similar to those normally found in library records. Recent changes in
the cataloging rules will cause some discrepancies between headings used here and
those used in libraries. There are no "see" or "see also" references since each country's
list is relatively short.
Entomology is of major importance in almost every field of agriculture. Since it is
not possible to list all serial publications of interest to the entomologist, the titles
included have been largely limited to those concerned directly with entomology as evi-
denced by words in the title or agency name. The few exceptions are serials which deal
with the related topics of general zoology, plus animal and plant pathology.
Since 1966 there has been a cooperative acquisition arrangement between the Uni-
versity of Florida Hume Agricultural Library and the Florida Department of Agricul-
ture and Consumer Services Division of Plant Industry Library (DPI Library). Mater-
ials relating to entomological taxonomy are acquired by the DPI Library, while all other
publications in entomology normally are acquired by Hume Library. Many serial titles
have been acquired for Hume Library and for the DPI Library through the assistance
of the Florida Entomological Society as exchanges for their publication, Florida En-
tomologist. This practice should be continued, and additional titles on this list should
be acquired in this fashion for the appropriate area libraries. Titles in this list marked
King: Latin Entomological Serials 31
by a dot are presently held by one or both of these libraries.
Many people have assisted in the compilation of this list. In order to enlarge and
correct this list, the Florida Entomological Society sponsored a mailout of the draft to
104 scientists and research agencies throughout Latin America. There were fourteen
responses. Several were extensive lists compiled locally, and they have contributed
immeasurably to this list. We appreciate the opinions expressed as well as the additional
titles suggested. Special thanks are due to Dr. Nelson Papavero of Brasil and Dr.
Marion Elgueta of Chile for their contributions.
June Jacobson, as Librarian of the DPI Library, has verified holdings for her collec-
tion and added specialized titles not otherwise located. The University of Florida Latin
American Collection has been a source of much information. Its Head Librarian, Rosa
Q. Mesa, as author of Latin American Serial Documents, has furnished many relevant
titles. Many thanks go to those who have translated and corrected in both Spanish and
Portuguese. I would personally like to thank the Florida Entomological Society and its
officers for their encouragement and for their financial support of the survey of Latin
American scientists. I wish to thank the SHARE Office, Institute of Food and Agricul-
tural Sciences, University of Florida, for financial support and to express my thanks to
Tomas Zoebish for searching these references. Any omissions and errors are entirely
Most important, thanks are due to Dr. Thomas J. Walker, University of Florida
Department of Entomology. This list was initiated as a direct result of his strong in-
terest in the literature of Latin America. We hope this final list will be of use to other
scientists and librarians who have an interest in the entomology of Latin America.
Florida Agricultural Experiment Station Journal Series No. 6925.
July 31, 1985 Ann H. King
Esta lista de casi 300 obras cubre, comprensivamente, la literature entomol6gica de
latinoam6rica. Tambien se incluyen titulos en las areas de zoologia general y en
patologia vegetal y animal cuando estos, por su contenido, son esencialmente en-
tomol6gicos. Las obras incluidas han sido publicadas por agencies gubernamentales a
various niveles, al igual que por compafiias privadas y sociedades cientificas.
America Latina ha sido siempre un area de gran interns para los cientificos en la
Universidad de Florida. Consecuentemente, la adquisici6n de publicaciones cientificas
de esta Area ha tenido siempre alta prioridad. Esta lista de publicaciones en entomologia
se ha compilado con el prop6sito de poder identificar titulos para su possible adquisici6n.
En 1970, una lista significativa titulada The Serial Literature of Entomology: A
Descriptive Study, fue compilada por Gloria M. Hammack, y publicada bajo los auspicios
de la Sociedad Entomol6gica de America y la National Science Foundation. En 1968,
una lista similar mas no publicada fue compilada localmente por Jane Rayborn de la
Division de Industria Agricola de Florida. La present lista fue iniciada con el prop6sito
de combinar ambas secciones, la de Rayborn y la de Hammack, y a su vez incorporar
titulos aparecidos a partir de 1970.
Esta lista incluye titulos publicados por agencies gubernamentales a various niveles
al igual que aqu6llos publicados por compahias commercials y sociedades cientificas.
La agrupaci6n de estos titulos se ha hecho primero por Areas geograficas (Cariba,
Am6rica Central) y luego por paises de origen. La cita bibliografica se encuentra
32 Florida Entomologist 69(1) March, 1986
generalmente bajo el titulo, except cuando la publicaci6n es producida por una
sociedad, agencia de gobierno o universidad, en cuyo caso generalmente se usa el
nombre de dicha agencia. Este tipo de cita es similar al normalmente encontrado en los
catalogos de las bibliotecas. Los cambios recientes en las reglas de catalogaci6n podran
causar discrepancies entire las citas aqui listadas y aqu6llas usadas en las bibliotecas.
No se ha hecho referencia a "v6ase" o vasee tambien," ya que cada pais tiene una lista
La entomologia es de gran importancia en casi todos los campos de la agriculture.
Ya que no es possible compilar una lista de todas las obras que puedan ser de interns
para el entom6logo, los titulos aqui incluidos se han limitado a aqu6llos que eviden-
temente, palabras en el titulo o nombre de la agensia son relevantes para entom6logos.
Como excepgi6n se han incluido publicaciones que tratan con t6picos relacionados, esto
es, zoologia, y patologia animal y vegetal.
Desde 1966 ha existido un acuerdo de adquisici6n cooperative entire la Biblioteca
Agricola (Hume) de la Universidad de la Florida y la Biblioteca de la Divisi6n de la
Industria Agricola de Florida (DPI Library). Todo material relacionado con taxonomia
entomol6gica es adquirido por la DPI Library, mientras que otras publicaciones en-
tomologicas son normalmente adquiridas por Hume. Muchas obras han sido adquiridas
para Hume y para DPI Library a trav6s de la Sociedad Entomol6gica de la Florida, a
cambio de su publicaci6n, Florida Entomologist. Esta practice debe de continuarse, y
las obras adicionales en esta lista deberAn ser adquiridas y designadas de este modo a
la biblioteca apropiada. Los titulos en esta lista que estan marcados con un "dot" se
encuentran presentemente en una o ambas bibliotecas.
Muchas personas han asistido en la compilaci6n de esta lista. Con el prop6sito de
expandir y corregir esta lista, la Sociedad Entomol6gica de la Florida envi6 un for-
mulario por correo a 104 cientificos y agencies de investigaci6n en latinoam6rica. Re-
cibieron catorce respuestas. Algunas fueron listas extensas compiladas localmente, y las
cualer contribuyeron inmensamente a esta lista. Estamos muy agradecidos por las
sugerencias dadas al igual que por las obras propuestas. Estamos especialmente ag-
radecidos al Dr. Nelson Papavero del Brasil y al Dr. Marion Elqueta de Chile por sus
La sefora June Jacobson, en su capacidad como bibliotecaria del DPI Library, ver-
ific6 las posesiones en su colecci6n y asisti6 en la adici6n de obras especializadas, las
cuales no se hubieran podido localizar de otra manera. La Colecci6n LatinoAmericana
de la Universidad de la Florida ha sido una fuente de much informaci6n. Su Bib-
liotecaria, la Sra. Rosa Q. Mesa, autora de Latin American Serial Documents, ha
proveido muchos titulos relevantes. Muchas gracias les damos a todos los que ayudaron
en las traducciones y correcciones en ambos espafol y portugues. Personalmente le
quiero dar las gracias a la Sociedad Entomol6gica de la Florida y a sus dirigentes por
el estimulo y las asistencias econ6micas en reconocimiento de los cientificos
LatinoAmericanos. Yo aquisiera agradecer a la oficina SHARE, del IFAS, Universidad
de Florida, por la ayuda financiera otorgada para esta investigation. Tambien quisiera
dar las gracias a Tomas Zoebish por la investigation de estas referencias. Cualquier
error u omisi6n son completamente mios.
Un agradecimiento muy especial se le debe al Dr. Thomas J. Walker del Depar-
tamento de Entomologia de la Universidad de la Florida. Esta lista fue iniciada como
resultado por su gran interns en la literature de latinoamerica. Esperamos que esta lista
sea fitil para otros cientificos y bibliotecarios interesados en la entomologia de
31 de julio 1985 Ann H. King
King: Latin Entomological Serials 33
Esta lista de aproximadamente 300 revistas cientificas cobre de completamente a
literature entomol6gica da America Latina. Inclui igualmente revistas nas areas de
zoologia geral, patologia animal e vegetal, quando tem substantial conteldo en-
tomol6gico. Ao revistas incluidas sao publicadas quer por agnncias governamentais a
vArios niveis, quer por firmas comerciais e sociedades cientificas.
A America Latina, ter sido sempre uma Area de grande interesse para os cientistas
da Universidade da Fl6rida. Consequentemente, a aquisicao de publicao5es desta Area
tem recebido, sempre, alta prioridade. A present lista de peri6dicos entomol6gicos esta
sendo publicada como um meio de identificar publicac6es para possivel aquisicao.
Em 1970, Gloria M. Hammack compilou uma significant relacao relevant entitulada
The Serial Literature of Entomology: a Descriptive Study, sob o patricinio da Sociedade
Entomol6gica da America e da National Science Foundation. Esta ralacao continha um
nimero substantial de titulos oriundos da America Latina. Em 1968, uma listagem
semelhante (nao publicada) foi produzida localmente por Jane Rayborn, da Biblioteca
do Departamento de Agricultura e Servigo para Consumidores da Fl6rida, Divisao de
Indastria Agricola (DPI Library). Esta lista foi feita pela necessidade de combinar as
seccqes relevantes das listas de Hammack e Rayborn, e incorporar publicag6s iniciadas
A present lista inclui titulos publicados por ag6ncias governamentais a varios niveis,
bem como por editors comeciais e sociedades cientificas. 0 seu arranjo foi feito por
Areas geograficas principals (i.e. America Central), e, seguidamente, por pais de origem.
A ordem de entrada 6 feita por titulo, except quando a publicagao 6 editada por uma
associagao ag&ncia, ou universidade, caso em que se usa geralmente a ag6ncia pat-
rocinadora. Este tipo de citaqgo 6 id6ntico ao normalmente utilizado mos catalogos de
bibliotecas. AlteravOs recentes das normas de catalogaaio poderao causar algumas dis-
crepancias entire os titulos usados aqui e os usadas nas bibliotecas. Uma vez que a lista
referente a cada pais 6 relativamente curta, refernncias tais como "veja" ou "veja tam-
b6m" sao omitidas.
A entomologia 6 de importAncia primordial em quase todos os campos da agriculture.
Nao send possivel listar todas as revistas de interesse para os entomologistas, in-
cluimos aqui apenas os titulos directamente ligados A entomologia, usando como crit6rio
o nome da revista ou da ag6ncia editor. Incluem-se, tamben publicaq6es nas Areas
relacionadas de zoologia geral e patologia animal ou vegetal.
Desde 1966, existe um acordo de aquisicao cooperative entire a Biblioteca Agricola
(Hume) da Universidade da Fl6rida e a Biblioteca da Divisiao de Indfstria Agricola
(DPI Library). Materials relacionados com a sistematica entomol6gica sao adquiridos
pela DPI Library, sendo todas as outras publicag6es normalmente adquiridas pela Hume
Library. Muitas das publicaqAos t&m sido adquiridas pela a Hume Library e para DPI
Library por permuta com a Florida Entomologist, publicacao official da Sociedade En-
tomol6gica da America. Esta forma de actuavaoao 6 desejavel e novos titulos constantes
desta lista deveriam ser adquiridos do mesmo modo para as bibliotecas das respectivas
Areas. Titulos que nesta lista vao marcados cor um "dot" existed presentemente numa
ou em ambas as bibliotecas.
Muitas pessoas participaram na compilagao desta lista. Para aumentar e corrigir esta
lista, a Sociedade de Entomol6gica de F16rida financiou o envio de c6pias preliminares
a 104 cientistas ou instituigbe selecionadas em toda a America Latina. Foram obtidas
catorze respostas. Varias delas eram listas exaustivas, compiladas localmente, que con-
34 Florida Entomologist 69(1) March, 1986
tribuiram imensamente para a present lista. Aqui agradecemos as opini6es expresses
e as sugest6e adicionais. Agradecemos especialmente ao Dr. Nelson Papavero do Brasil
e ao Dr. Marion Elqueta do Chile pelas suas contribuigoes.
June Jacobson, como Bibliotecaria da DPI Library, verificou as existencias na re-
spectiva colecvao e acrescentou titulos da especialidade que nao haviam sido pr6-
viamente localizados. A Colecdo Latinoamericana da Universidade de Fl6rida, foi igual-
mente uma fonte valiosa de informaa&o. A sua bibliotecaria chefe, Rosa Q. Mesa, autora
dos Latin American Serial Documents, forneceu muitos titulos relevantes. Muitos ag-
radecimentos soa dirigidos aos tradutores para o espanhol e portugues. Gostaria de
pessoalmente, agradecer a Sociedade de Entomol6gica de Fl6rida e respective direccao
pelo seu encorajamento e suporte financeiro no levantamento dos cientistas latino-
americanos. Gostaria de agradecer ao departamento SHARE, do IFAS, Universidade
da Fl6rida, por apoio financeiro a esta investigaqao. Os meus agradecimentos vao tam-
bem para Tomas Zoebish, peia pesquisa bibliogrAfica realizada. Por quaisquer omissbes
e enganos sou inteiramente responsavel.
Agradecimentos especiais sao devidos a Dr. Thomas J. Walker do Departamento de
Entomol6gica da Universidade de Fl6rida. Esta lista foi iniciada como resultado director
do seu grande interesse pela literature da America Latina. Esperamos que esta lista
final seja dtil a outros cientistas e bibliotecarios que tambem tenham interesse na en-
tomologia da America Latina.
31 julio 1985 Ann H. King
LIST OF BIBLIOGRAPHIES SEARCHED
Aliaga de Vizcarra, Irma. Guia de Publicaciones Periodicas Agricolas y Conexas de
Bolivia. Sociedad de Ingenieros Agronomos de Bolivia. Boletin Bibliografico,
Arboleda-Sepilveda, Orlando. Directorio de las Publicaciones Periodicas de la Biblioteca
Conmemorativa Orton. Institute Interamericano de Ciencias Agricolas. 1966-68.
Volume with Suplo. 1-5.
Badillo, V. M. and C. Bonfanti. Indice Bibliografico Agricola de Venezuela. Fundaci6n
Eugenio Mendoza, 1957.
Bibliografia Brasileira de Zoologia, v. 5, 1970; v. 6-7, 1971-72; v. 11, 1975-77; v. 12,
1977-78; v. 13, 1978-79.
Catalogue of the Imperial College of Tropical Agriculture, University of the West In-
dies, Trinidad. Periodicals Section, p. 631-725. G. K. Hall, 1975.
Gropp, Arthur E. A Bibliography of Latin American Bibliographies Published in Period-
icals. Scarecrow, 1976.
Hammack, Gloria M. The Serial Literature of Entomology: a Descriptive Study. En-
tomological Society of America, 1970.
HernAndez de Caldas, Angela. Publicaciones Periodicas Bioagricolas Latinoamericanas:
un Directorio. Universidad de Narifio, Instituto Tecnol6gico Agricola, 1966.
Irregular Serials and Annuals. R. R. Bowker, 5th ed., 1978-79, 6th ed. 1980-81.
Levi, Nadia. Guia de Publicaciones Peri6dicas de Universidades Latinoamericanas. Uni-
versidad Nacional Aut6noma de M6xico, 1967.
MArquez, Orfila and Belkys GutiBrrez. Guia de Publicaciones Periodicas Agricolas de
Venezuela. Repiblica de Venezuela, Ministerio de Agric'ltura y Cria, Oficina de
Comunicaciones Agricolas, 1972.
Mesa, Rosa Q. Latin American Serial Documents. R. R. Bowker.
v. 1, Colombia, 1968 v. 10, Peru, 1973
v. 2, Brasil, 1968 v. 11, Uruguay, 1973
King: Latin Entomological Serials 35
v. 3, Cuba, 1969 v. 12, Venezuela, 1977
v. 4, Mexico, 1970 Costa Rica, unpublished
v. 5, Argentina, 1971 El Salvador, unpublished
v. 6, Boliva, 1972 Guatemala, unpublished
v. 7, Chile, 1973 Haiti, unpublished
v. 8, Ecuador, 1973 Nicaragua, unpublished
v. 9, Paraguay, 1973 Panama, unpublished
New Serial Titles 1950-70: Subject Guide. R. R. Bowker, 1975.
New Serial Titles 1971-80: Subject Guide. R. R. Bowker, 1971-80.
Pan American Union. Index to Latin American Periodical Literature. G. K. Hall.
Peri6dicos Brasileiros de Cultura. Edicao Preliminar. Institute Brasileiro de Biblio-
grafia e Documentavao. 1956.
Peri6dicos Brasileiros de Cultura. Institute Brasileiro de Bibliografia e Documentacao.
Rayborn, Jane. World List of Entomological Literature. Florida Division of Plant Indus-
try, unpublished, 1968.
Shelby, Charmion, ed. Latin American Periodicals Currently Received in the Library
of Congress and in the Library of the Department of Agriculture. The Library
of Congress, 1944.
Ulrich's International Periodicals Directory. 18th ed., 1979-80, 19th ed., 1980, and 20th
Velasquez, Pablo and Ramon Nadurille. Catilogo Colectivo de Publicaciones Peri6dicas
Existentes en Bibliotecas de la Republica M6xicana. v. 1-2. Institute Nacional
de Investigaciones Agricolas, 1968.
Academia de Ciencias de Cuba. Institute de Zoologia. Informe Cientifico-Tkcnico.
(Habana. Direcci6n de Publicaciones de la A.C.C.) no. 1, 1977-
Academia de Ciencias de Cuba. Institute de Zoologia. MiscelAnea Zool6gica. (Habana)
no. 1, 1975-
CatAlogo de la Fauna Cubana. Serie 4: Ciencias Biol6gicas. (Habana. Centro de Informa-
ci6n Cientifica y Thcnia, Universidad de la Habana) no. 35, 1975-
Control de Plagas. (Habana) v. 13, no. 1, 1950-
Cuba. Secretaria de Agricultura, Industria y Trabajo. Secci6n de Sanidad Vegetal.
Anuario. (Habana) 1926/27-
Cuba. Secretaria de Agricultura, Industria y Trabajo. Secci6n de Sanidad Vegetal.
Boletin. (Habana) no. 1, 1917-no.4, 1923?
Cuba. Secretaria de Agricultura, Industria y Trabajo. Secci6n de Sanidad Vegetal.
Bulletin. [In English] (Habana) no. 1, 1917.
Cuba. Secretaria de Agricultura, Industria y Trabajo. Secci6n de Sanidad Vegetal.
Circular. (Habana) no. 1, 1916-no.6, 19?
Cuba. Secretaria de Agricultura, Industria y Trabajo. Secci6n de Sanidad Vegetal.
Informe de los trabajos. (Habana) 1924/25?
Poeyana. Serie A. no. 1, 1964-
Poeyana. Serie B. no. 1, 1964-
Revista Kuba de Medicina Tropical y Parasitologia. (Habana) 1947- [Was: Revista de
Parasiologia, Clinica y Laboratorio. (Habana) 1935-36; Revista de Medicina Trop-
ical y Parasitologia, Bacteriologia, Clinica y Laboratorio. (Habana) 1936-46; Re-
vista de Medicina Tropical y Parasitologia. (Habana) 1945-46.]
Florida Entomologist 69(1)
Moca. Estaci6n Nacional Agron6mica. Publicaci6n. Serie E. Entomologia y Zoologia.
(Santo Domingo) v. 1, 1927-
[no. 1 as Laboratorio de Entomologia. Boletin. (Santo Domingo) no. 2 as
Laboratorio de Entomologia. Circular. (Santo Domingo)]
Moca. Estaci6n Nacional Agron6mica. Laboratorio de Entomologia. Informe de la Sec-
ci6n de Entomologia.
Nouvelles Agronomiques. (Antilles-Guyane)
Revue Agricole: Organe du Service de l'Agriculture de la Guadeloupe et Dependances.
(Basse-Terre) v. 1-3, 1926-28?
Jamaica. Department of Agriculture. Entomological Bulletin. (Kingston) 1921-32.
Jamaica. Department of Agriculture. Entomology Circular. (Kingston) 1921-34.
Journal of Agriculture of the University of Puerto Rico. (Rio Piedras) v.1, 1917-
School of Tropical Medicine of Puerto Rico. Report. (San Juan) 1943-47?
University of Puerto Rico. Estaci6n Experimental Agricola. Boletin.
TRINIDAD AND TOBAGO
Caribbean Plant Protection Commission. Quarterly Report (Port of Spain, Food and
Agriculture Organization of the U.N.)
Commonwealth Institute of Biological Control. Quarterly Report. (Trinidad)
Tropical Agriculture. (University of the West Indies) v.1, 1924-
El Agricultor Costarricense. (San Jose) 1943-
* Brenesia. (San Jos6, Museo Nacional de Costa Rica) v.1, 1972-
Ciencias Veterinarias. (Heredia, Universidad Nacional) v.1, 1980-
Revista de Biologia. (San Jos6, Universidad de Costa Rica) v.1, 1953-
* Turrialba. (San Jos) v.1, 1953-
El Salvador. Centro Nacional de Tecnologia Agropecuaria. Boletia Thcnico. no.1, 1970?-
El Salvador. Centro Nacional de Tecnologia Agropecuaria. Publicaciones. Varias v.1,
El Salvador. Direcci6n General de Investigaciones Agron6micas. Secci6n de En-
tomologia. Boletin Tecnico. (San Salvador) v.1, 1960-
Entomologia Economica Hondurefa. Boletin T6cnico. no.6, 1958-
Abejas y miel. Boletin. (Mexico, D.F. Direcci6n General de Avicultura y Otras Especies
Menores) v.1, 1964-
Chapingo, MWxico. Escuela Nacional de Agricultura. Departamento de Parasitologia.
Memoria del Dia del Parasit6logo. v.1, 1963-
Enciclopedia Apicola. (Mexico, D.F. Editorial Agricola Mexicana) v.1, 1953-
King: Latin Entomological Serials 37
* Fit6filo. (San Jacinto) afo 1, 1942-
* Fitopatologia Mexicana. (Chapingo, Escuela Nacional de Agricultura, Laboratorio de
Fitopatologia, Sociedad M6xicana de Fitopatologia. v.1, 1962-?)
* Folia Entomol6gica Mexicana. (Mexico, D.F., Sociedad Mexicana de Entomologia)
El Informador Apicola. (M6rida) 1964-
* Institute de Salubridad y Enfermedades Tropicales. Revista. 1939-65.
* MWxico (City). Universidad Nacional. Institute de Biologia. Serie Zoologfa. (M6xico,
D.F.) v.39, 1967- [Supercedes, in part, and continues v. nos. of its Anales.]
M6xico. Comisi6n de Parasitologia Agricola. Boletin. no.l, 1900-no.4, 1908.
M6xico. Comisi6n de Parasitologia Agricola. Circular. no.l, 1903-no.75, 1908.
M6xico. Comisi6n de Parasitologia Agricola. Las Plagas de la Agricultura. Entrega
no.l, 1902-no.12?, 1903.
Mexico. Comisi6n Nacional para la Erradicaci6n del Paludismo. CNEP Boletin. (M6xico,
D.F.) no.l, 1957-
Mexico. Consejo Superior de Salubridad. Boletin Extraordinario. Documentos Oficiales
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M6xico. Institute de Higiene. Secci6n de Parasitologia. Monografias. no.l, 1923-no.4,
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Florida Entomologist 69(1)
* Archivo de Ciencias Biol6gicas y Naturales, Te6ricas y Aplicadas. (Buenos Aires,
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Argentine Republic. Institute de Sanidad Vegetal. Boletin. (Buenos Aires).
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Argentine Republic. Institute de Sanidad Vegetal. Laboratorio Central de
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Boletin Fitosanitario. (Buenos Aires, Secretaria de Estado de Agricultura y Ganaderia
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Buenos Aires. Direcci6n de Lucha contra las Plagas. Circular D. 1957-
Buenos Aires. Institute Cientifico de Medicina Veterinaria. Boletin Tecnico. 1957-
* Buenos Aires. Institute de Entomologia Sanitaria. Publicaciones del Instituto Reg-
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Buenos Aires. Institute de Patologia Vegetal. Hoja Informativa. 1966?-
Buenos Aires. Institute de Patologia Vegetal. Publicaciones. Serie A. 1945-54.
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* Buenos Aires. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia." Com-
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King: Latin Entomological Serials 39
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[Title varies: Memoria.]
Comit6 Interamericano Permanente Antiacridiano. Reuni6n Anual. (Buenos Aires)
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Revista Argentina de Entomologia. (Buenos Aires) v.1, 1935-v.2, 1944.
Revista de Investigaciones Agropecuarias. Serie 4: Patologia Animal. (Buenos Aires)
* Revista de Investigaciones Agropecuarias. Serie 5: Patologia Vegetal. (Buenos
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* Universidad de Buenos Aires. Facultad de Agronomia y Veterinaria. Revista de la
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Universidad Nacional. Facultad de Agronomia y Veterinaria. Institute de Parasitologia
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* Universidad Nacional. Museo. Anales. Secci6n Zoologica. (La Plata) no.l, 1893-no.3,
Universidad Nacional. Museo. Anales. Nueva Serie. Zoologia. (La Plata) no.l, 1953-
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Bolivia. Departamento de Sanidad Vegetal. Boletin Entomol6gico. (La Paz) no.1, 1951-
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* Acta AmazOnica. (Manaus, Instituto Nacional de Pesquisas da Amazonia) v.1, 1971-
Acta Biol6gica Paranaense. (Curitiba, Universidade Federal do Parana) v.1, 1972- [was
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Florida Entomologist 69(1)
Apicultor. (Niteroi, Associagao Fluminense de Apicultores) v.1, 1961-
Apicultor. (Porto Alegre, Associacao Gaucha de Apicultores "AGA") v.1, 1965-
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Arquivos de Entomologia. Serie A. (Pelotas, Instituto Agron6mico do Sul, Escola de
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* Ciencia e Cultura. (Sao Paulo, Sociedade Brasileira para o Progresso da Ci6ncia) v.1,
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* Dusenia: Publicaqdo Periodica de Scientia Naturali. (Curitiba) 1950-
Entomologia Medica. (Sao Paulo) v.1, 1962-v.3, 1965.
Entomologista Brasileiro. (Sao Paulo) Ano 1, 1908-ano 3, 1910. [was: Entomologo]
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Fertilizantes, insecticides e races. F.I.R. (Sao Paulo) v.1, 1959?-
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Fitossanidade. (Fortaleza. Grafica Recol.) no.l, 1974-
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Iheringia Zool6gica. (Rio Grande do Sul, Museu Rio-Grandense de Ciencia Naturais.
Department de Ciencia e Cultura da Sec.) 1957-
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Instituto de Biol6gico e Pesquisas Tecnol6gicas. Boletim. (Curitiba) v.1, 1941-
Instituto de Medicina Tropical de Sao Paulo. Revista. 1958-
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Museu Nacional. Boletin. Nova s6rie Zool6gia. Boletim. no.l, 1942-
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Natura. (Bahia, Universidade Federal da Bahia. Institute de Biologia) 1975-
Res6menes de Malariologia. (Rio de Janeiro) 1942-46?
Resumos de Malariologia e Doengas Tropicais. (Rio de Janeiro) 1948-
King: Latin Entomological Serials 41
* Reuniao de fitossanitaristas do Brasil. Anais. 1st, 1956? -
Revista Brasileira de Apicultura. (Rio de Janeiro) v.1, 1922-v.1, 1923.
* Revista Brasileira de Biologia. (Rio de Janeiro, Academic Brasileira de Ci6ncias) v.1,
* Revista Brasileira de Entomologia. (Sao Paulo, Sociedade Brasileira de Entomologia)
* Revista Brasileira de Malariologia. (Rio de Janeiro, Servico Nacional de Malaria) v.1,
* Revista Brasileira de Malariologia e Doenvas Tropicais. (Rio de Janeiro. Dept. Na-
cional de Endemias Rurais) v.1, 1949-
* Revista Brasileira de Pesquisas Medicas Biol6gicas/Brazilian Journal of Medical and
Biological Research. (Sao Paulo, Editora M6dico) v.1, 1968-
* Revista de Agricultura (Piracicaba) v.1, 1926-
* Revista de Entomologia. (Rio de Janeiro. Thomaz Borgmeier, O.F.M.) v.1, 1931-
no.22, 1951. Supplemento, no.l, 1941.
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1939. [became: Studia Entomol6gica]
Rio Grande do Sul, Brasil (State) SecretAria da Agricultura, Indfstria, e Comercio.
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Sao Paulo. Brasil (State) Departamento da Producao Animal. Serie de Vulgarizagao.
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Sociedade Brasileira de Entomologia. Anais de ReuniAo Anual. (Sao Paulo. Escola
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* Sociedade Entomol6gica do Brasil. Anais. (Rio de Janeiro) v.1, 1972-
Sociedade Entomol6gica do Brasil. Boletim. (Rio de Janeiro) No.l, 1922-no.6, 1923.
* Studia Entomologica, Revista Internacional de Entomologia. First Series. (Rio de
Janeiro) No.l, 1952-no.3, 1955.
* Studia Entomologica, Revista Internacional de Entomologia. New Series. (Rio de
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* Summa Phytopathol6gica. (Sao Paulo. Grupo Paulista de Fitopatologia, Sociedade
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* Universidade de Minas Gerais. Museu de Historia Natural. Boletim. Zoologia. (Belo
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Universidade Federal do Parana. Boletim. Zoologia. (Curitiba) v.2, 1965-v.5, 1972. [be-
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Boletin Agricola Shell. (Santiago)
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Boletin de Sanidad Vegetal. (Santiago de Chile) v.1, 1941-v.3, 1943.
Chile. Comisi6n de Parasitologia Agricola. Circular. (Santiago de Chile) 1903-1908.
Florida Entomologist 69(1)
Chile. Departamento de Sanidad Vegetal. Boletin Tecnico. (Santiago de Chile) no.l,
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Chile. Estaci6n de Patologia Vegetal. Publicaciones. (Santiago de Chile?) no.l, 1900-
Chile. Estaci6n Nacional Agron6mica. Laboratorio de Entomologia. Boletin. (Santo
Chile. Estaci6n Nacional Agron6mica. Laboratorio de Entomologia. Circular. (Santo
Chile. Servicio Nacional de Salubridad. Departamento de Parasitologia. Memoria annual.
(Santiago de Chile) 1952?-
Colecci6n insects portadores de enfermedades. (Santiago de Chile. Sub. Depto. de
Educaci6n Sanitaria) 1956-
Instituto de la Patagonia. Anales. (Punta Arenas) v., 1970-
* Investigaciones Zool6gicas Chilenas. (Santiago, Centro de Investigaciones
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versidad de Chile) v.1, 1951-
Revista Chilena de Historia Natural Pura y Aplicada. (Valparaiso, Museo de Historia
Natural) v.1, 1897-v.55, 1960/63. [title varies: Revista Chilena de Historia Natu-
* Santiago de Chile. Museo Nacional de Historia Natural. Boletin. t.1, 1908-
Sociedad Entomol6gica de Chile. Boletin. (Santiago) no.l, 1928-no.2, 1929.
Sociedad Entomol6gica de Chile. Boletin Semestral. (Santiago) no.l, 1973-no.2, 1975.
Universidad de Chile. Biblioteca Tecnica de Parasitologia. Contribuci6n a la Bibliografia
'Chilena de Parasitologia. (Santiago) v.1, 1954-
Universidad de Chile. Centro de Estudios Entomol6gicos. (Santiago) Publicaciones.
no.l, 1960-no.10, 1970. [became: Universidad de Chile. Departamento de
Biologia. Publicaciones Entomol6gicas]
Universidad de Chile. Departamento de Biologia. Publicaciones Entomol6gicas. (San-
tiago) v.ll, 1974- [was: Universidad de Chile. Centro de Estudios Entomol6gicos.
Universidad de Chile. Departamento de Parasitologia. Boletin de Informaci6n Tecnica.
(Santiago) v.1, 1945?- v.7, 1952. [title varies: Boletin de Informaci6nes
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Universidad de Chile. Departamento de Parasitologia. Boletin de Informaciones
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Universidad de Chile. Departamento de Parasitologia. Publicaciones Instructivas de
Caracter Sanitario. (Santiago)
Universidad de Chile. Departamento de Parasitologia y Servicio Nacional de Salud.
Indice Bibliografico Parasitol6gico de Articulos de Revistas. (Santiago) 1952-
Universidad de Chile. Facultad de Agronomia. Estaci6n Experimental Agron6mica.
Boletin T4cnico. (Santiago) v.1, -
Actualidades Biol6gicas. (Medellin, Universidad de Antioquia) 1972-
* Agriculture Tropical. (Bogota, Asociaci6n Colombiana de Ingenieros Agr6nomos) v.1,
* Caldesia. (Bogota, Instituto de Ciencias Naturales) v.1, 1940-
Colombia. Comisi6n Central para la Extinci6n de la Langosta. Boletin de los trabajos.
King: Latin Entomological Serials 43
no.l, 1912-no.20, 1913?.
Revista Colombiana de Entomologia. (Sociedad Colombiana de Entomologia) v.1,
Sociedad Biol6gica de Bogota. Anales. (Bogota) v.1, 1945-
Sociedad Colombiana de Entomologia. Memorias. 1976?-
Sociedad Colombiana de Entomologia. Resumenes. 1976?-
Universidad Nacional de Colombia. Facultad de Medicina Veterinaria y de Zootecnia.
Revista. (Bogota) 1928-
Revista Ecuatoriana de Entomologia y Parasitologia. (Quito. Centro Ecuatoriano de
Investigaciones Entomol6gicas) v.1, 1953-
Sanidad Vegetal. (Quito, Ministerio de Economia) no.1, ?
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varies: 1905-1907? as Instituto Nacional de Bacteriologia.]
Paraguay. Ministerio de Agricultura. Servicio de Extensi6n Agricola y Ganaderia. Bole-
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Universidad Nacional. Institute de Ciencias Basicas. Informes Cientificos. (Asunci6n)
Archivos de Biologia Andina. (Lima, Universidad Nacional Mayor de San Marcos. In-
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Fitopatologia. (Lima, Asociaci6n Latinoamericana de Fitopatologia) v.1, 1966-
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forme General de la Campana Peri. 1964/65?-
Lima, Peru. Estaci6n Experimental Agricola. Informe Mensual. 1927-
Peri. Departamento de Malaria. Publicaciones. (Lima, Direcci6n General de Salud Pib-
* Peril. Servicio de Investigaci6n y Promoci6n Agraria. Informe especial. no.l, 1962-
Peri. Servicio Nacional de Erradicaci6n de la Malaria. Boletin. (Lima) no.l, 1960-
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no.l, 195?- [Title varies: Informe Anual de Actividades.]
* Revista Peruana de Entomologia Agricola. (Lima, Sociedad Entomol6gica Agricola
del Peril) v.1, 1958-v.2, 1959.
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Centrum voor Landbouwkundig Onderzoek in Surinam.
Landbouwproefstation in Surinam. Bulletin
* De Surinam Landbouw. v.1, 1953-
44 Florida Entomologist 69(1) March, 1986
Apicultor Americano. (Montevideo) v.1, 1954-v.2, 1958.
Liminas Entomol6gicas. (Montevideo)
* Museo Nacional de Historia Natural. Comunicaciones Zool6gicas. (Montevideo) v.1,
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* Sociedad Uruguaya de Entomologia. Revista. v.1, 1956-
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(Montevideo) no.l, 1920-no.5, 1924.
Uruguay. Comisi6n Central de Defensa Agricola. Defense Agricola. Memoria. (Mon-
Uruguay. Comisi6n Central de Defensa Agricola. Defense Agricola. Publicaci6n. no.l,
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Agriculture Tropical. (Maracay) v.1, 1951-
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Boletin de Entomologia Venezolana. (Caracas, Ministerio de Sanidad y Asistencia So-
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* Caracas. Museo de Ciencias Naturales. Boletin. t.1, 1955-
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no.l, 1982 -
Maracay, Venezuela. (State) Instituto Nacional de Agricultura. Boletin Tkcnico.
(Maracay) v.1, 1956-v.6, 1951.
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tomologia y Zoologia. Circulares.
Maracay, Venezuela. (State) Instituto Nacional de Agricultura. Division de
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Revista de Medicina Veterinaria y Parasitologia. (Maracay. Facultad de Medicina Vet-
erinaria, Universidad Central de Venezuela) v.1, 1939-
Sociedad de Ciencias Naturales. Memoria. (La Salle, Caracas) v.9, 1949?-
Sociedad Venezolana de Ciencias Naturales. Boletin. (Caracas) v.1, 1931-
Tijeretazos sobre Malaria. (Caracas, Divisi6n de Malariologia) v.1, 1938-?
Universidad Central. Facultad de Agronomia. Revista. (Maracay) v.1, 1952-
* Universidad de Zulia. Facultad de Agronomia. Revista. (Maracaibo) v.1, 1960?-
Venezuela. Direcci6n de Agricultura. Division de Entomologia y Zoologia. Boletin T6c-
nico. (Caracas) no.l, 1949?-
Venezuela. Direcci6n de Agricultura. Division de Fitopatologia. Boletin. (Caracas) no.1,
Venezuela. Direcci6n de Agricultura. Division de Sanidad Vegetal. Boletines. (Caracas)
Venezuela. Direcci6n de Agricultura. Division de Sanidad Vegetal. Circulares.
Venezuela. Direcci6n de Ganaderia. Compafia de Sanidad Animal.
Venezuela. Direcci6n de Malariologia y Saneamiento Ambiental. Boletin Informativo.
(Maracay) v.1, 1961-
Venezuela. Direcci6n de Malariologia y Saneamiento Ambiental. Informe annual.
Venezuela. Direcci6n de Malariologia y Saneamiento Ambiental. Reuniones Quin-
King: Latin Entomological Serials 45
cenales: Informe. (Caracas) 1961?-
Venezuela. Direcci6n de Malariologia y Saneamiento Ambiental. Reuniones Quin-
cenales: Minuta. (Caracas) 1961?-
Venezuela. Division de Malariologia. Boletin Informativo. (Maracay) no.l, 1959-?; 2
series. v.1, no.l; 1961-
Venezuela. Division de Malariologia. Publicaciones. (Caracas) no.l, 1938-
Venezuela. Institute Nacional de Agricultura. Division de Entomologia y Zoologia.
Venezuela. Institute Nacional de Agricultura. Division de Fitopatologia. Boletin.
Venezuela. Institute Nacional de Agricultura. Division de Fitopatologia. Noticias
Florida Entomologist 69(1)
STOCHASTIC POLYPHENISM: COPING WITH UNCERTAINTY
THOMAS J. WALKER*
"A potentially important source of individual phenotypic variation is examined
which has heretofore received little recognition as a general and probably wide-
spread phenomenon." (Cooper and Kaplan 1982, p. 135)
Polyphenism (i.e., polymorphism in which non-genetic differences cause the develop-
ment of the contrasting forms) can be conditional or stochastic. In conditional
polyphenism, a genotype responds to different current environments that predict differ-
ent future environments by producing different, appropriate phenotypes. For example,
short days may cause the development of the diapause phenotype, appropriate to
winter, and long days may cause the development of the nondiapause phenotype, appro-
priate to summer. In stochastic polyphenism, a genotype responds to differences in its
environment that occur with probabilities approximating the probabilities of different
future environments. For example, if 30% of winters are severe enough to require
diapause and the other 70% yield a corresponding advantage to nondiapausing morphs,
a genotype would produce the diapause phenotype 30% of the time and the nondiapause
phenotype the other 70%. Modeling and empirical evidence support the concept of
In a constant environment natural selection should lead to genotypes that develop
into individuals adapted to that environment. In a predictably varying environment
natural selection should operate to yield genotypes that cause the development of indi-
viduals appropriate to whatever environment is coming next. In fact, genotypes that
anticipate the deterioration of a local habitat or the progression of the seasons are
commonplace in insects. Such genotypes switch development into one path or another
in response to present conditions that predict the future.
In environments that vary unpredictably-i.e., in truly uncertain environments-
the outcome of evolution is hard to predict. The thesis of this paper is that genotypes
which have probabilistic or stochastic output should prosper in uncertain environments
and that such genotypes exist in insects.
Although I arrived at the notion of stochastic genes independently, the concept is
not original. D. A. Roff (1975) investigated models in which the probability of dispersal
was a function of genotype. W. S. Cooper and R. H. Kaplan (1982, p. 136; also, Kaplan
and Cooper 1984) made a strong case for "intra-genotypic strategy-mixing or, less for-
mally, adaptive coin flipping."
If genotypes have stochastic outputs, outputs of a single genotype could vary con-
tinuously (e.g., intensity of diapause, minimum duration of flight in a dispersing indi-
vidual) or discontinuously (e.g., diapause vs. nondiapause; winged vs. apterous).
Stochastic outputs of both types are indicated, but only discontinuous variation will be
discussed here because it is more striking and easier to treat.
*Thomas J. Walker is a Professor of Entomology in the Department of Entomology and Nematology, University
of Florida. His research deals mostly with the systematics, acoustic behavior, and ecology of crickets and katydids,
but he also studies butterfly migration. Current address: Department of Entomology and Nematology, University
of Florida, Gainesville, Florida 32611.
Insect Behaviorial Ecology-'85 Walker 47
A GENETIC POLYMORPHISM C CONDITIONAL POLYPHENISM
G (E, or Ep or Es) p 1
G2 (E or E2 or Es) P2 2
D STOCHASTIC POLYPHENISM
B POLYPHENISM P,
P N2 (00-y)%) P2
0 -, C2) Es= y % N, + (loo-y) % N2
Es y% P1
(3) G 1 __
S (1 00-y)% P2
Fig. 1. Diagrams illustrating different types of polymorphism. (A) In genetic
polymorphism, different phenotypes (P1 and P2) develop as a result of differences in
genotype (G1 and G2). (B) In polyphenism, different phenotypes develop as a result of
nongenetic differences (N1 and N2). (C) In conditional polyphenism, the nongenetic
differences are environmental circumstances (El and E2) that predict different future
environments to which the different phenotypes are adapted. (D) In stochastic
polyphenism, (1) the nongenetic differences occur at probabilities that correlate with
the probabilities of future environments to which the different phenotypes are adapted;
(2) these stochastically occurring alternatives can be considered to comprise an "environ-
ment" (Es), and (3) this environment causes the two phenotypes to be produced in the
proportions of N1 and Ng.
Discontinuous variation occurring within a deme between individuals of the same
ontogenetic stage is called polymorphism (see Kennedy 1961). Polymorphism can be
divided into genetic polymorphism and polyphenism depending on what sort of differ-
ences lead to the development of the distinctive phenotypes. In genetic polymorphism
the distinctive phenotypes result from differences in genotype, whereas in polyphenism
nongenetic differences are responsible (Fig. la,b). It should be noted that polyphenism
depends on a genotype that can be switched by the environment from one path of
development to another.
Two modes of selection may lead to polyphenism. First, and most obvious, is that
natural selection will increase genotypes that respond to different environments by
producing different, appropriate phenotypes.' Such genotypes are responsible for condi-
tional polyphenism (Fig. Ic). The defining aspect of this type of polyphenism is that
48 Florida Entomologist 69(1) March, 1986
0 I- 1 --- 100
I modified by uncontrolled
S variation in environment
and/or in genotype
of P, %) of P2 (%)
I ideal conditional
low E1 threshold E2 high
Environmental Variable (E)
Fig. 2. The concept of threshold as applied to conditional polyphenism. An environ-
mental variable ("E", abscissa) that varies continuously is divided into two regimes (E1
and E2) by a threshold. The transition from 100% of one phenotype (P1) to 100% of the
other (P2) is likely to be sigmoid rather than rectilinear, and the best estimate of
threshold becomes the value of E at which 50% of the exposed population develop into
P1 and 50% into P2.
development of a phenotype adapted to a specific future environment is conditional upon
a particular present environment that is correlated with the future one. Examples
include development of seasonally appropriate diapausing or nondiapausing individuals
(Beck 1980) or flight-worthy or flightless individuals (Harrison 1980) in response to
photoperiod. Conditional polyphenism depends on a genotype responding to one en-
vironmental condition by developing one phenotype and to an alternative condition by
developing another. The transition between these two conditions is a threshold that
delimits the production of 100% of one phenotype from the production of 100% of the
other (Fig. 2). Ideally the threshold is sharp, but in practice it is likely to be blurred,
because individuals in the responding population are not genetically identical (and there-
fore vary in their thresholds) and because individuals in a population do not experience
identical environments even under carefully controlled circumstances (Fig. 2).
The second mode of selection is that a genotype may increase because it produces
alternative phenotypes in proportions that approximate the probabilities of future envi-
ronments that favor the respective phenotypes. Such selection would lead to stochastic
polyphenism (Fig. Id).
If stochastic polyphenism occurs, (1) genes must be able to effect a stochastic output
and (2) such genes must be able to increase at the expense of genes that have a deter-
ministic output. If these two conditions are met, (3) one would expect to find examples
of stochastic polyphenism in insects living in environments exhibiting important, short-
to medium-term, unpredictable variation. The next three sections of this paper deal
with these subjects.
Insect Behaviorial Ecology-'85 Walker 49
CAN GENES GAMBLE?
For genes to produce a stochastic output they must make use of environmental
differences that occur in the right proportions without pattern. To use the metaphor of
Cooper and Kaplan (1982), they must flip coins (properly biased to produce the needed
proportions of heads and tails).
Recent advances in genetics suggest a variety of ways that genes might, in effect,
gamble with the phenotype of their carrier. Transposable genetic elements ("jumping
genes"), identified by Barbara McClintock in maize and now known to be commonplace
in other plants and in animals (Fedoroff 1984), offer one such possibility. The develop-
mental effect of a mobile element depends on its position in the genome. For instance,
if it attaches to site "A" 40% of the time and to site "B"-60% of the time, the outcome
would be 40% of one phenotype and 60% of another. The genomes would be the same
except for the positions of their components, but the phenotypes produced would be
determined stochastically.2 Another possibility was suggested by Spudich and Koshland
(1976), who demonstrated individual variation in genetically identical bacterial cells
grown in a homogeneous environment. In trying to account for the variations, they
noted that certain molecules occur in such small numbers in cells that their numbers
are subject to "Poissonian variation" when cells divide and that this could lead to signif-
icant random variation in phenotype.
A final example is from immunology. The ability to produce millions of different
antibodies is passed from generation to generation in a limited number of genes. These
genes specify a "kit of [genetic] components," which are "shuffled" in the developing B
lymphocytes leading "to a different result in each of millions of lines of cells" (Leder
1982, p. 102). Even millions of different phenotypes can be produced stochastically by
a single genotype.
CAN STOCHASTIC GENES COMPETE?
If stochastic genes can arise by mutation from deterministic forebears, how well will
they compete. Simulation modeling is a suitable way to investigate this question. Using
a microcomputer, I developed two models that test stochastic genes in uncertain envi-
ronments." In each model, the environment was made to vary without pattern between
The first model concerned insects that sometimes benefit from diapause and some-
times benefit by continuing development. One environmental state (e.g., mild winter),
occurring p% of the time, was more favorable to the nondiapause phenotype. The other
state (e.g., severe winter), occurring (100-p)% of the time, was more favorable to the
diapause phenotype. Diapause was determined by three alleles at one locus: DD, produc-
ing the diapause phenotype; DN, nondiapause phenotype; and Ds, y% diapause
phenotype and (100-y)% nondiapause phenotype.5 The following conditions were
specified to start the model: dominance relations of the alleles, initial frequencies of
alleles, proportion of generations that environment favors diapause phenotype (i.e., p),
proportion of diapause phenotypes produced by Ds (i.e., y), relative fitness of diapause
phenotype when the environment is less favorable to it, relative fitness of nondiapause
phenotype when the environment is less favorable to it, number of generations per run
of model, and number of runs.
Table 1 gives representative results for the first model. When allele Ds was recessive
to the other two alleles, it quickly replaced them under a variety of scenarios--even if
the probability of the diapause phenotype deviated greatly from the probability of the
diapause-favoring environment (e.g., Table 1, A and B). When the other alleles were
recessive, they persisted more than 400 generation under a wide range of assumptions,
Florida Entomologist 69(1)
00 011t-4- ooor--
00 010OCO 0OC1~''
o~ dd 66oo
W C 3
0 C0 D
A A A
>0 000 0000CD
00 -100 000
66 666 6666
Insect Behaviorial Ecology-'85 Walker
0 ^ h ;
a a ^ ^
Q <<: H <
_ h tf
0 O <
2 Mh f
S tf ?
^ ^ ^ f
g f d
'c d -
E in c
o~r oo t- oC
0a c0-Oar oc
O- 00 0-0 0
0 00000 0C
oo ooooo os
66 66666 6e
08 00 r0fl 0-
oo coooo oe
00 00L--4 0 <
66 66666 6e
0 0 0 0
0 0 1 0
0 0 00 0
0 000 0
52 Florida Entomologist 69(1) March, 1986
and the average frequency of Ds after 400 generations depended on the assumptions.
For example, when y was made to deviate from p and the witnesses of the disfavored
phenotypes were kept equal, the frequency attained by Ds decreased (Table 1, C, lines
2 and 3). On the other hand, if the witnesses of the disfavored phenotypes were made
unequal, the frequency realized by Ds was increased by changing its "setpoint" (i.e., y)
toward a higher proportion of the less disfavored phenotype (Table 1, D).
The second model concerned insects in which a portion of diapausing individuals
remains dormant for more than a year (e.g., Prebble 1941, Powell 1974, Ushatinskaya
1976). In the wheat-blossom midge (Sitodiplosis mosellana), for example, diapause may
last as long as 12 years; the emergence from year to year is irregular, but the trend is
for a smaller percentage of a cohort to emerge each succeeding year (Barnes 1952). A
presumed advantage of prolonged diapause is that the individual may skip a poor or
disastrous annual growing season and emerge in a favorable one. If bad seasons cannot
be foretold by the time diapause is broken", the individual should, on average, benefit
by breaking diapause the first year-because the probability of happening on a good
year is the same each year but the mortality associated with staying in diapause accumu-
lates year after year. On the other hand, a gene that caused most carriers to emerge
the first year and lesser numbers to emerge subsequent years should be superior to
any single-output gene. The model to test the superiority of genes with mixed outputs
made certain assumptions7; and, again, initial conditions had to be set."
Table 2 gives representative results. When recessive, the allele with mixed output
(Ps) sometimes quickly replaced the other alleles (Table 2, A-C). When unfavorable
growing seasons were rare relative to the annual attrition rate, Ps was less successful
(Table 2, B). Similarly, when unfavorable growing seasons were less severely unfavor-
able, Ps attained at lower frequencies (Table 2, C).
The results from these two models suggest that alleles producing a stochastic output
can increase in frequency at the expense of competing deterministic alleles and some-
times completely replace them.
Other workers have tested models of stochastic alleles. Roff (1975) created a variety
of complex models in which genes controlled the probability of dispersal among subpopu-
lations that varied independently and randomly in their finite rates of increase (gener-
ally between 0 and 2.8). In his simplest version, he assumed the population was fixed
for an allele that determined a stated dispersal probability, and he showed that popula-
tions with probabilities intermediate between 0 and 1.0 would persist. In another ver-
sion, dispersal was controlled by a single locus with two alleles-a non-dispersal allele
and a dispersal allele that produced dispersal and non-dispersal phenotypes in a specified
ratio. The probability that dispersers would survive was also specified. Roff found that
when the stochastic allele was recessive it either reached 100% and the population
survived or, in cases where low survival of dispersers had been specified, the non-dis-
persal allele reached 100% and extinction ensued. When he made the stochastic allele
partially dominant, the model reached an enduring genetic polymorphism under a wide
range of assumptions about the probabilities that the two genotypes permitting disper-
sal would produce dispersal phenotypes.
Cooper and Kaplan (1982) were first to focus directly upon the fitness of genotypes
with stochastic output. They used "decision trees" to compare the witnesses of genotypes
with deterministic and stochastic outputs in uncertain environments and concluded that
genotypes with mixed output would win over deterministic genotypes under a variety
of conditions and even if their output did not match exactly the probabilities of the
All efforts at simulation modeling of genotypes with mixed and pure outputs in
uncertain environments have yielded similar results: stochastic alleles can replace or
reach a lasting genetic polymorphism with alleles that have a single output.
Insect Behaviorial Ecology-'85 Walker 53
Because stochastic polyphenism is expected to coexist with conditional polyphenism
and genetic polymorphism, with all three affecting the same traits, clearcut examples
may be difficult to identify even if the phenomenon is frequent. Some examples from
insects are instructive.
Aphids are remarkably adapted to the study of polyphenism. All species are
polymorphic and produce several to many generations of viviparous parthenogenetic
females each year (Dixon 1985a). Parthenogenetic reproduction in aphids apparently
includes no recombination, making all parthenogenetically produced females genetically
identical to their mothers, except for new mutations (Blackman 1979, Tomiuk and
Wohrmann 1982, Dixon 1985b).9 In other words, the asexual descendants of any parth-
enogenetic female (and there may be millions) constitute a clone. The abundant
polymorphism within parthenogenetic aphid lineages is thus entirely polyphenism (Lees
1961, Dixon 1985b).
Much of aphid polyphenism is clearly conditional. For example, the production of
sexual females, which lay winter-hardy eggs, is cued variously by short photoperiods,
cool temperatures, and season-related changes in host plants (Dixon 1985a). Winter is
More relevant to possible stochastic determination of morphs is the production of
winged and wingless midseason morphs (alatae and apterae). The relative witnesses of
these morphs depend on the future suitability of the home host plant and the quantity
and quality of other host plants, reachable only by flight. Host quality and availability
are less easily predicted than is winter. In most cases that have been studied, production
of alatae is only to a degree conditional to environment correlates of decreasing home
host quality (e.g., crowded conditions and decline in nutritional quality of the host)
(Dixon 1985a). An important aspect of the response to these correlates is that rarely is
it 100%; the response is ordinarily 20 to 95% alatae (and 80 to 5% apterae). The switch
is thus generally between development of one phenotype (apterae) and development of
two phenotypes (apterae and alatae) rather than between one phenotype and another.
Shaw's (1970a) studies of the bean aphid (Aphis fabae) are representative. When he
reared offspring of apterae under the crowded conditions of 200/bean stipule, about 40%
developed into alatae (65%, if the mothers had been crowded too). This degree of crowd-
ing could be construed to be the approximate threshold for the switch from 100% apterae
to 100% alatae (see Fig. 2), except that densities nearly half as great and densities twice
as great still produced a mixture of phenotypes (Fig. 3).
Most other studies have yielded similar results. Lees (1967) found that crowding
caused most apterous female vetch aphids (Megoura viciae) to produce mixtures of
alatae and apterae, generally 70 to 95% alatae. Sutherland (1969) reported that crowding
pea aphids (Acyrthosiphon pisum) resulted in a maximum of 80 to 85% alatae (in green
and pink strains, respectively). Lamb and MacKay (1979) tested ca. 500 clones of pea
aphids taken from alfalfa fields in southern Ontario and found that crowding increased
production of alatae to an average of 66% (but never to 100%). On the other hand, Watt
and Dixon (1981) reported 100% alatae when one strain of the English grain aphid
(Sitobion avenae) was crowded-though another strain of the same species gave rise to
few alatae when treated the same way.
The general lack of a 100% response in production of alatae suggests that selection
has favored genotypes that produce a mixture of alatae and apterae under conditions
signaling a decline of the home host. The proportions of the mix can be influenced by
genotype, as indicated by consistent interclone variability in proportions of alate and
54 Florida Entomologist 69(1) March, 1986
Aphis fabae ,c -
(%) / [%J
100- -- ------
I r I I
0 100 200 300 400
Population Density (larvae/stipule)
Fig. 3. Response of bean aphids to crowding (data from Shaw 1970a). Filled points
and solid line are for the crowded progeny of uncrowded apterae. Open points and upper
dashed line are for crowded progeny of apterae that were themselves crowded 100-400
per stipule. No alatae are produced at densities below 100 larvae/stipule. The difference
between the two data sets shows that the environment of the mother influences the
likelihood of her offspring being alatae. (The lower dashed line is for crowded offspring
of alatae. Offspring of alatae are never alate, again demonstrating a maternal influence.)
(Lines are eye-fitted.)
apterous pea aphids (Lamb and MacKay 1979). Mixed-output genotypes and genetic
differences, as well as conditional responses, apparently contribute to wing dimorphism
in midseason aphids.
Dispersal polymorphisms in sexually reproducing insects are common, and they sel-
dom are attributable to simple genetic differences, although the proportions of the
morphs can be altered by selection (Harrison 1980). A case in point is wing dimorphism
in field crickets (Gryllus spp.).
Field crickets are either long-winged or short-winged (Walker and Sivinski 1986).
The short-winged morph, like the apterous aphid, never flies. The long-winged morph,
characterized by the metathoracic wings extending well beyond the tegmina, has, like
an alate aphid, the external equipment for flight. (And again like the alate aphid, it does
not necessarily fly-Shaw 1970b; Walker, unpublished data.)
Collections of Grylus rubens the most abundant field cricket in southeastern
United States, vary greatly in the proportions of long- and short-winged morphs
(Veazey et al. 1976). When progeny of field-collected females are reared under con-
trolled conditions or in outdoor cages, they are generally 5-95% long-winged (Walker,
unpublished data). Walker (unpublished data), starting with field-collected stock,
selected for long and short wings under controlled temperature and photoperiod
(25+ 10C, 16L:8D). The two strains diverged until one was ca. 95% long-winged and the
other was ca. 95% short-winged. Even though selection was 100%, in neither strain was
the rejected phenotype eliminated in seven generations (Fig. 4). The change in propor-
Insect Behaviorial Ecology-'85 Walker 55
1 2 3 4 5 6 7
Fig. 4. Results of 100% selection for the long- and short-winged morphs of a field
cricket (Gryllus rubens) for seven generations. Long- and short-winged parents were
collected in Gainesville, Fla., March 1983, and all subsequent rearing and selection was
at 25+1C, 16L:8D. Only long-winged morphs were used as parents in the L x L line
and only short-winged morphs were used in the S x S line.
tions of morphs showed that some of the original polymorphism was attributable to
genetic differences (as in other Gryllus: Harrison 1979, Roff 1984). Conditional
polyphenism is also part of the explanation, as shown by a variety of studies of related
species by others (see Alexander 1968) and as shown by exposing the F6 generation of
the selected G. rubens lines to three rearing conditions (Fig. 5). However, the persistent
dimorphism, in the face of selection and under the almost constant environment of a
controlled temperature room, suggests that G. rubens genotypes have been selected to
maintain a mixed output-i.e., that stochastic polyphenism is also involved in the
The pitcher-plant mosquito Wyeomyia smithii is dimorphic in its warm season larval
development. Some third instars enter diapause and others continue development.
Using standardized rearing conditions, including a 15-h photoperiod, C. A. Istock and
co-workers studied environmental and genetic determinants of this dimorphism. In one
series of experiments, Istock et al. (1975), varied the amount of food per larva. Diapause
incidence dropped to as low as 15% with ample food and increased to 100% (among
survivors) when food was deficient. In another series of experiments, Istock et al.
(1976) maintained conditions that gave 56% diapause in the unselected stock and selected
for diapause and nondiapause strains. The course of selection resembled that in Fig. 4
in that progress was irregular and at the end of the study both strains were still
producing both phenotypes. After 15 generations of selection for fast, nondiapause
development, diapause incidence was 4%. After 7 generations of selection for diapause,
56 Florida Entomologist 69(1) March, 1986
Conditions L x L (F6) S x S (F6)
16L:8D 98% LW 6%LW
VBC diet GLW Gs
1/cup 2%SW 94%SW
16L:8D 89% LW 3%LW
Cricket Chow GL G
LW ~ SW -
50/jar 11%SW 97%SW
1 1L:13D 47%LW 1%LW
Cricket Chow G Gs
Fig. 5. Response of two strains of Gryllus rubens to three sets of rearing conditions
at 25 1C. Each strain was in its sixth generation of 100% selection for long or for short
wings (see Fig. 4). GLW = genotype(s) of the L x L strain; Gsw = genotype(s) of the
S x S strain; LW = long-winged phenotype; SW = short-winged phenotype. Rearing
conditions differed in photoperiod, diet, type of container, and number of crickets per
container. (In the language of Fig. 1, each set of rearing conditions is an "Es".) (For
treatments of L x L, n = 47, 165, and 97; for S x S, n = 63, 170, 86; the proportion of
LW from GLW is significantly lower for the short-day treatment than for the two long-
incidence was 88%. In contrast to these results at a 15-h photoperiod, under fall photo-
periods (14 h or less) all third instars diapause. Diapause dimorphism in W. smithii,
like wing dimorphism in G. rubens, includes genetic polymorphism and conditional
polyphenism as well as what seems to be stochastic polyphenism.
J. G. Sternburg and G. P. Waldbauer (see Waldbauer 1978) found that the silk moth
Hyalophora cecropia has two modes of emergence in central Illinois. About 10% of the
moths emerge in late May (Group I) and the other 90% in late June (Group II). The
adult moths are too short-lived to survive the gap between modes, suggesting reproduc-
tively isolated populations. However, the majority of the progeny of Group I moths
emerge the following year in Group II and a minority of the progeny of Group II emerge
in Group I the following year-proving that Group I and Group II mergers belong to
the same Mendelian population. Four generations of selection for Group I emergence
changed the proportion emerging in Group I to ca. 80% (Waldbauer 1978). The propor-
tion of the wild population that emerged in Group I during the years of selection varied
between 4.7% and 10.0%. That genotypic differences are partly responsible for the
bimodal emergence is demonstrated by the pronounced increase in proportion of Group
I mergers in the selected lineage. That conditional polyphenism is slightly involved is
implied by the year to year fluctuation in the proportion of Group I mergers in the
wild population (but that fluctuation could also result from changes in genotype frequen-
Insect Behaviorial Ecology-'85 Walker 57
cies from one year's generation to the next). Finally, the failure of four generations of
selection to establish a pure line of Group I mergers shows that mixed emergence is
entrenched in H. cecropia genotypes, as expected in stochastic polyphenism.
If stochastic polyphenism should be a frequent evolutionary response to uncertainty,
why might it go generally unrecognized? One reason is that it occurs in combination
with other types of polymorphism, and a researcher who proves that one of the other
types occurs is likely to emphasize what is explained by accepted principles rather than
what is not. A related reason is that chance seems unsatisfactory as a scientific explana-
tion. This misses the point that selection may favor events that are stochastic in effect
even though deterministic in means (a flipped coin obeys the laws of physics, but the
effect is close enough to 0.50 probability that football games are started with one). A
third reason is that in stochastic polyphenism individual fitness is subordinate to fitness
at another level. Those who have been trained that selection at the individual level is
the only safe way to view evolution may be reluctant to accept as an adaptation a
phenomenon that causes some individuals to have reduced fitness for the long term
advantage of stochastic genes. However, as Dawkins (1982) has emphasized, selection
is as much (or more) the differential survival of genes as the differential reproduction
How have those who eschewed stochastic polyphenism explained the type of data
used here to support it? Most have ignored the issue. Some have cited maternal influ-
ence. A mother aphid that can make her offspring be y% one phenotype and (100-y)%
another phenotype may, on average, have more grandprogeny than one that produces
100% of what is, on average, the fittest phenotype. Maternal influence seems unlikely
when the development of phenotypic differences occurs long after maternal contact has
ceased (e.g., wings in field crickets). More to the point, maternal influence is not actually
an alternative to stochastic polyphenism. The mother's genotype (rather than the pro-
geny's) simply becomes the one that flips the coin that determines the progeny's
phenotype. Furthermore, in aphids, the mother's and progeny's genotypes are generally
the same! For sexually reproducing insects, a more credible alternative to stochastic
polyphenism is a form of genetic polymorphism-viz., polygenic inheritance with
threshold effect (Fig. 6) (see Falconer 1981, Chap. 18).10 In the case of Gryllus rubens
or Wyeomyia smithii, the production of both phenotypes after 7-15 generations of 100%
selection could be attributed to a polygenic complex's normally slow response to selec-
tion (Falconer 1981, Chap. 12)." Finally, frequency dependent selection should be men-
tioned. Although it is a common cause of genetic polymorphisms (Clarke 1979, Rausher
1986), it cannot account for polymorphism within aphid clones or for the difficulty in
artificially selecting for pure breeding strains of long-winged crickets, warm-season
diapausing mosquitoes, or early emerging moths.12
When consequences of maladaptation are unequal, what should be the effect on the
setpoint of stochastic polyphenism? In most of Table 1 the disadvantage of diapause in
a nondiapause environment was made equal to the disadvantage of nondiapause in a
diapause environment (A, B, C). If the disadvantages are made unequal (as in Table 1,
D), stochastic alleles that produce more of the least disadvantaged phenotype achieve
higher frequencies.13 If one disadvantage is eliminated, then the optimal setpoint be-
comes 100% of the phenotype that is never disadvantaged-i.e., the polymorphism
What more general evolutionary principles is stochastic polyphenism a subset of?
One is that natural selection does not always favor genotypes that produce higher
average numbers of offspring. Gillespie (1977) showed that genotypes with lesser vari-
Florida Entomologist 69(1)
A / "t I/ / /0//
C 95% 5%
Low Threshold High
--- scale of "underlying continuous variable" --
Fig. 6. Diagram illustrating polygenic inheritance with threshold effect as an alternative
explanation of wing dimorphism in G. rubens (Fig. 4). An underlying continuous variable
controls the dimorphism (e.g., the concentration of a wing-promoting substance). This
variable is controlled by genes at many loci with additive effects, producing a normal
distribution of levels. Some of the loci are linked in opposite phase, prolonging the time
required to develop a pure breeding short- or long-winged strain. A. Unselected. B.
After 6-8 generations of 100% selection for long-winged morph. C. After 6-8 generations
of 100% selection for short-winged morph. (Diagrams modified from Falconer 1981).
Insect Behaviorial Ecology-'85 Walker 59
ances in offspring numbers can increase at the expense of those with higher average
numbers of offspring. In the case of stochastic temporal fluctuations of the environment,
he found the best measure of fitness to be the geometric mean of offspring number,
averaged over time (the procedure used by Cooper and Kaplan, 1982, in their decision
tree analyses). Another principle is that evolution is more easily understood when
analyzed at the level of the replicator (Dawkins 1982). As Cooper and Kaplan (1982, p.
145) explained, a strategy-mixing genotype causes some individuals to be stuck with an
inferior phenotype-"for the sake of the long term advantage of the genotype."
What terms should be used for the types of polyphenism? Dawkins (1980), writing
of evolutionarily stable strategies, distinguished conditional from mixed or stochastic
strategies in individual behavior. This distinction aided my theorizing about
polyphenism. Cooper and Kaplan (1982; also, Kaplan and Cooper 1984) used predictive
and coin-flipping to name the same distinction and were first to apply it to "adaptive
phenotypic plasticity," which includes polyphenism. Both sets of terms are appealing
and nearly self-defining. I chose Dawkins' because they are more likely to be familiar
to behavioral ecologists. Less this choice be viewed as deemphasizing Cooper and Kap-
lan's priority in applying the distinction to polyphenism, I end as I began by quoting
from their synopsis (1982, p. 135): "The variation is a product of the action of genetically
controlled stochastic processes; metaphorically, it is produced because individuals are
genetically programmed to 'flip coins' to decide what characteristics to adopt. Thus it
is not the variable phenotypic traits themselves that are genetically specified, but only
the nature of the coin-flipping process that will ultimately determine them. ... coin-flip-
ping strategies of this kind are robust and can evolve under a variety of conditions."
I thank James B. Kring for help with the aphid literature, Chai-Lin Tan for a
summary of the literature on prolonged diapause, and H. M. Wallbrunn for guiding me
to Wright's studies of inbred guinea pigs; I am grateful to J. E. Lloyd, Todd Pickard,
John Sivinski, Frank Slansky, Sue Wineriter, and Tony Zera for constructive criticism
of the manuscript. The study of wing polymorphism in G. rubens was aided by NSF
grant BNS 81-03554. Florida Agricultural Journal Series No. 7065.
'In sexually reproducing animals, natural selection increases genotypes indirectly-
by altering frequencies of alleles that in turn alter the probability of particular
genotypes forming in the next generation and by altering linkage relationships.
2The effect of a transposable element is that of a mutation and its back mutation
being exceedingly frequent, making the occurrence of the phenotypes they produce
unpredictable except on a probabilistic basis. That a mobile element can actually control
a polymorphism has been demonstrated in the bacterium Salmonella (Simon et al. 1980,
p. 1370), in which the inversion of a 970-base-pair DNA sequence "behaves like a flip-flop
switch activating and inactivating H2 gene transcription. Depending on the frequency
of switching, a fraction of the population expresses one flagellar antigen while the rest
of the cells express the other flagellar antigen."
3These models were written in BASICA for an IBM PC. They are menu-driven and
suitable for classroom use. I will make copies for anyone sending me two blank diskettes
(one of which I will return with the programs).
4This was accomplished through the random-number generating function of
5Phenotypes were assigned individuals in the proportions y:(100-y). For large popu-
lations this procedure should not differ in effect from true stochastic assignment with
probabilities y and (100-y).
60 Florida Entomologist 69(1) March, 1986
6In some instances, circumstances in one growing season foretell circumstances the
following year, making possible control of prolonged diapause by conditional
polyphenism. For example, a dense population (as in an outbreak) can predict scarce
resources or high parasitoid populations for the next year.
7The assumptions were: (1) Bad years occur at random. (2) Three alleles control
duration of diapause-Po, which causes emergence after one winter; PT emergence
after two winters; and Ps, 50% emergence after one winter, 25% emergence after two
winters, 12.5% emergence after three winters, etc. up to w winters, with all remaining
individuals emerging after the wth winter (e.g., for w=4, 12.5% emerged the last
winter). (3) The annual mortality caused by remaining in diapause is a constant propor-
tion of survivors. (4) Each year's cohort (i.e., those entering their initial winter of
diapause) is numerically independent of the number that had broken diapause earlier
that year to produce it but dependent on whether the growing season had been good
'In running the prolonged-diapause model these values were set: dominance relations
of alleles, initial frequencies of alleles, maximum number of winters in diapause for
individuals controlled by Ps, probability of a bad growing season, number going into
diapause after a bad growing season (as a proportion of the number for a good growing
season), number of generations per run, number of runs.
9Mutation-caused morphs should be easily separable from polyphenism because such
morphs would occur rarely and, once present, the responsible allele should be passed
to all parthenogenetically produced descendents of the original mutant individual.
'A way to refute the hypothesis of polygenic inheritance with threshold effect as an
alternative explanation (for data that suggest stochastic polyphenism in sexually repro-
ducing insects) is the establishment of homozygous strains through inbreeding. As
strains become more and more homozygous through generations of inbreeding, the
polymorphism may be maintained if it is stochastic polyphenism and should be reduced
and then eliminated if it is dependent on genetic difference. As an example of this
approach, Wright and Chase (1936) (see also, Festing 1976) studied white spotting in
inbred and outbred lines of guinea pigs and found that in inbred lines the degree of
spotting was almost entirely under nongenetic control. Individuals ranging from nearly
white to nearly "self colored" (i.e., 5-95% white spotted) occurred in the same litter. In
one inbred line the environment controlled 97% of the variance, and less than 10% of
this environmental component was attributable to age of mother and other factors
common to litter mates-89% of the variance was random. Even in an outbred
line, 52% of the variance in degree of spotting was random.
"One of the causes of slow response is linkage of polygenes of opposite sign. If such
linkage is tight, the group of linked loci can act as a "supergene" for stochastic
polyphenism-by keeping the level of the "underlying continuous variable" close enough
to the threshold of response for individuals of both phenotypes to occur.
'2Frequency dependent selection occurs, by definition, whenever fitness of a
genotype is an inverse function of its frequency. Wing dimorphism in crickets is partly
a result of genetic differences among individuals, and the frequency of alleles for long-
and short-winged morphs may depend in part on long-winged and short-winged morphs
becoming more fit on average as they (respectively) become rarer. On the other hand
the relative success of long- and short-winged morphs must depend in part on uncertain
meteorological events. Frequency dependent selection may occur as a result of unpre-
dictable environmental changes but it is not an adaptation to them.
lCooper and Kaplan (1982) derived this formula for calculating the optimal setpoint
for a stochastic genotype:
qo- X F1 + Po
X1,0 Ko,o K'l, 0h,1
where qo = proportion of phenotype 0
Insect Behaviorial Ecology-'85 Walker
K,\o = finite rate of increase, in environment favoring phenotype 1, of phenotype 0
-o,o = finite rate of increase, in environment favoring phenotype 0, of phenotype 0
P, = probability of environment favoring phenotype 1
proportions of phenotypes 0 and 1 are qo and 1-qo.
Applying this formula to the circumstances in Table 1,C, yields an optimal y of 0.704;
to the circumstances in Table 1,D, an optimal y of 0.995.
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Insect Behaviorial Ecology-'85 Rausher 63
COMPETITION, FREQUENCY-DEPENDENT SELECTION,
AND DIAPAUSE IN BATTUS PHILENOR BUTTERFLIES
MARK D. RAUSHER*
The pipevine swallowtail butterfly, Battus philenor, exhibits a diapause polymorph-
ism in east Texas. Approximately half the offspring of first-brood females enter and
remain in pupal diapause until the following year. The other half ecloses to form a
second brood. Evidence is presented to indicate that second-brood females compete for
oviposition sites. It is deduced that this competition causes the fitness of non-diapausers
to be frequency dependent: as the proportion of non-diapausers in the population in-
creases, their fitness decreases. A single-locus model showing that this type of fre-
quency dependence can maintain a stable diapause polymorphism is presented.
It is generally agreed that the availability of food plants greatly influences the
voltinism patterns of herbivorous insects (Opler and Langston 1968, Slansky 1974, Gil-
bert and Singer 1975, Shapiro 1975). In northern temperate communities, trees and
shrubs often produce a brief burst of high-quality, nutrient-rich foliage that rapidly
becomes a poor substrate for larval growth as nutrient levels decline and growth in-
hibitors increase in concentration (Feeny 1970, Raup and Denno 1983). In response to
this brief period of availability of high-quality larval food, many insects have evolved
to be univoltine and avoid having to use mature foliage. Other species, which feed on
mature foliage, may have a univoltine life cycle imposed upon them because slow growth
rates on low-quality foliage do not permit two generations to be completed during a
single growing season (Feeny 1970). By contrast, many forb-feeding insects, whose food
plants remain nutritious for much of the growing season, are multivoltine (Slansky
1974). Similarly, in areas of summer drought, where larval food plants rapidly senesce
with the onset of dry conditions, herbivorous insects tend to be univoltine. Multivoltine
species in these communities normally feed on host plants that remain green, such as
those growing along streams (Opler and Langston 1968, Shapiro 1975, Gilbert and
Many, if not most, populations of herbivorous insects are uniformly either single-
brooded (univoltine) or multiple-brooded (multivoltine). All individuals enter diapause
at approximately the same time of year and remain dormant for approximately the same
period of time (Danilevsky 1965, Danilevsky et al. 1970, Waldbauer 1978). This
monomorphic pattern is expected in species whose food supply remains constant through
much of the season (multivoltine species) or is present for a short pulse and then disap-
pears completely (univoltine species). Recently, however, it has become clear
that some species are polymorphic within a single population for the number of broods
that occur within a season (Helle 1968, Geyspitz 1968, Slansky 1974, Shapiro 1975,
Istock et al. 1975, 1976, Istock 1978, Waldbauer 1978).
While diapause polymorphisms have been well-documented, little is known about
*Mark D. Rausher is an Associate Professor in the Department of Zoology and the University Program in
Genetics at Duke University. In 1979 he received his Ph.D. under Paul Feeny at Cornell University. He has been
studying the behavioral ecology of Battus philenor butterflies since 1974. Other current research concerns the
ecological genetics of plant-insect interactions and modelling the evolution of searching behavior in herbivorous
insects. Department of Zoology, Duke University, Durham, North Carolina 27706.
Florida Entomologist 69(1)
the selective forces that preserve them in populations. Here I present evidence suggest-
ing that frequency-dependent selection, apparently mediated through competition for
oviposition sites, acts to maintain a diapause polymorphism in an east Texas population
of the pipevine swallowtail butterfly, Battus philenor.
PHENOLOGY AND DIAPAUSE IN B. philenor
In east Texas, populations of B. philenor overwinter in pupal diapause. In mid-
March, adults close to form the first brood (brood 1) and mated females begin laying
eggs on larval food plants. At this time of the year, most females search preferentially
for and oviposit on a small, perennial herb, Aristolochia reticulata (Aristolochiaceae).
In mid-May, some offspring of brood-1 adults close to form a second brood (brood 2).
Most females of this brood search preferentially for and oviposit on a closely related
host, Aristolochia serpentaria, because the foliage of A. reticulata has by this time
become tough and nutrient-poor (Rausher 1978, 1980, 1981, Rausher and Papaj 1983a).
Larval rearing experiments were performed in 1976 and 1977 to determine the
proportions of offspring of brood-1 and brood-2 adults that entered diapause.' In these
experiments, approximately half of the offspring of brood-1 individuals entered diapause
in both years (Table 1). There is thus clearly a polymorphism for diapause in the popu-
lation examined, but it is not known whether this phenotypic polymorphism is the result
of an underlying genetic polymorphism or whether the decision to diapause is based on
perception of a particular environmental cue2. Regardless of whether diapause is genet-
ically or environmentally determined, however, it seems reasonable to hypothesize that
natural selection actively maintains both diapausing and non-diapausing morphs within
the population studied.
In the remainder of this work I provide evidence that is consistent with this
hypothesis. In particular, I first document the existence of competition for oviposition
sites among brood-2 B. philenor females. I then describe why it is reasonable to infer
that the existence of such competition means that the fitness of the non-diapausing
morph, measured as representation in the next generation, depends on the frequency
of that morph in the population. Finally, I present a genetic model that shows how this
type of frequency-dependence can maintain the observed diapause polymorphism in
east Texas populations of B. philenor.
COMPETITION FOR OVIPOSITION SITES
By following female butterflies in the field, it is possible to determine the rate at
which they alight on (discover) host plants, the rate at which eggs are laid on host
plants, and the proportion of host plants alighted on that already bear eggs (Rausher
1979, 1983). Examination of temporal trends in these rates for brood-1 and brood-2
females in 1977 (the only year for which brood-2 data are available) provides the evi-
TABLE 1. FRACTION OF EXPERIMENTALLY-REARED PUPAE THAT DIAPAUSE.
Year Brood 1 Brood 2
1976 .41 (59)' .79 (24)
1977 .55 (60)
'Numbers in parentheses are sample sizes for diapausing and non-diapausing pupae.
Insect Behaviorial Ecology-'85 Rausher
dence that competition for oviposition sites occurs during the second brood.
During the first brood, oviposition rate shows no trend over time (Fig. la), probably
because females actively maintain a constant rate of oviposition (Rausher 1983). By
contrast, during the second brood, oviposition rate declines steadily (Fig. Ib). There are
EGGS LAID/10 MIN
I I I,
I I I I I I
li I ii l 1111 1 1l..l.,111 li ,l
5 10 15 20 25 30
Fig. 1 Temporal trend in oviposition rate (No. eggs laid/10 min.) over the course of
a brood. Each point represents pooled data for all individuals on the corresponding day.
Data based on a total of 1095 observation minutes for brood 1, 784 observation minutes
for brood 2. A. Brood 1. Spearman correlation coefficient (on pooled data), rs = .39,
NS. B. Brood 2. r, = -.72, P = .01.
Florida Entomologist 69(1)
three plausible explanations for this decline: (1) oviposition rate declines with female
age3, (2) alighting rates decline over time4, or (3) the number of acceptable host plants
decreases over time due to preemption by other females.5
It is unlikely that explanation 1 is correct. If aging of individuals were responsible
for the decline in oviposition rate during brood 2, then such a decline should also have
been observed in brood 1, which serves as a control for the observations made during
brood 2. Because a decline was not observed during brood 1 (Fig. la), the cause of the
decline during brood 2 is most likely due to some factor or process, unlike aging, that
was not operative during brood 1.
Analysis of alighting rates indicates that explanation 2 is also probably not correct.
Alighting rates do not decline over time during brood 2 (Fig. 2b; the increase in alighting
rates during brood 1(Fig. 2a) is for reasons explained in Rausher 1983), as would be
expected if this explanation were true. Instead, they remain more or less constant, and
the decline in oviposition rate occurs in spite of this constancy. While doubtless some
host plants were consumed by larvae during brood 2, the number is expected to be
small, since larval feeding activity in the habitat is minimal at this time (Rausher and
Feeny 1980). Evidently, so few were eaten that there was a negligible effect on host
abundances, and hence alighting rates.
By elimination, it would appear that explanation 3 is most likely the correct one.
Moreover, observed trends in proportion of alightings that are on host plants that
already bear eggs are precisely what are expected under this explanation. During the
second brood, there is a steady increase in the proportion of hosts that have previously-
laid eggs, from 0 at the beginning of the brood to approximately .85 by the end of the
brood (Fig. 3b)6. Consequently, the rate of alighting on potentially acceptable host
plants (plants without eggs) decreases markedly over the brood (Fig. 4b). Females
seem to be able to counteract this decrease in rate of alighting on plants without eggs
to some extent by increasing the probability of ovipositing as the brood progresses and
alighting rate falls (Fig. 5b). However, because even at the beginning of the brood the
probability of oviposition is high (i.e., approximately .6), the scope for such compensa-
tion is limited. The decline over time in rate of alighting on plants without previously
laid eggs is thus necessarily accompanied by a decline in oviposition rate.
The picture that emerges from this analysis, then, is as follows: females emerging
at the beginning of the second brood find that virtually all suitable host plants (primarily
A. serpentaria) are free of eggs. Oviposition rates at the beginning of the brood are
presumably limited by the rate at which host plants are discovered. Over time, as more
eggs are laid by females, a larger fraction of the host plants bear eggs or larvae. These
plants have been, in effect, preempted. By the end of brood 2, most host plants alighted
on have been preempted and only rarely does a female encounter a plant without eggs
or larvae. Consequently, oviposition rate, which is limited to a large extent by alighting
rate, has fallen to only about one quarter what it was at the beginning of the brood. It
is thus apparent that competition for oviposition sites is intense during the second
brood, even when only about half the offspring of brood-1 individuals emerge to form
the second brood.
Because it is technically impossible to manipulate with adequate controls the propor-
tions of the progeny of brood-1 individuals that eclose or enter diapause, it is not
possible to ascertain directly whether changing the frequency of individuals would affect
the mean fitness of non-diapausers. However, the data reported in the last section imply
that mean representational fitness of non-diapausers, defined as the number of offspring
represented in the next generation of adults (i.e., number of eggs laid times the average
Insect Behaviorial Ecology-'85 Rausher
PLANTS ALIGHTED ON/10 MIN
I I i
1 I a I I I I I I I I I I I I I I
I I I I 1 1 1 1 I 1 a I I a a I I I a I I I
Fig. 2 Temporal trend in alighting rate (No. host plants alighted on/10 min.) over
the course of a brood. A. Brood 1. r, = .98, P < .001. B. Brood 2. r, = .14, NS.
68 Florida Entomologist 69(1)
FRACTION OF PLANTS WITH EGGS
11 I 1 1 1 1 1**1 11 1
I I I I I I I I I I I I I I I I I I I
Fig. 3 Temporal trend in proportion of host plants alighted on that bear previously-
laid eggs. A. Brood 1. r, = .51, NS. B. Brood 2. r, = .74, P < .01.
Insect Behaviorial Ecology-'85 Rausher
PLANTS W/O EGGS/10 MIN
I I IL I l
LL I1IA I I l 111111111 I
I I I I I I I I I
I I a I I . a I
Fig. 4 Temporal trend in rate of alighting on plants without previously laid eggs. A.
Brood 1. r, = .95, P < .01. B. Brood 2. r, = -.71, P < .05.
. . . . I I I I
Florida Entomologist 69(1)
FRACTION OF PLANTS
S I I
I I I I .
& I I I I I I I I I I
I I I
Fig. 5 Temporal trend in probability of ovipositing, once alighting occurs, on host
plants without eggs. A. Brood 1. r, = -.85, P < .01. B. Brood 2. r, = .63, P < .05.
L * * a* I 1 1
Insect Behaviorial Ecology-'85 Rausher 71
probability of survivorship of offspring (Rausher 1985)), does in fact depend on their
frequency in the population.
Consider what would happen if all progeny of first-brood individuals emerged to
form the second brood. There would be approximately twice as many females competing
for the same limited number of oviposition sites. The 15% of the host plants that are
not typically preempted by the end of the second brood is not nearly sufficient to allow
twice as many females to lay the same mean number of eggs per capital with the same
average number of eggs laid per plant. One or more of several changes would therefore
occur: (1) Females might decrease the mean number of eggs laid per individual; (2)
Females might maintain the same mean fecundity by laying more of their eggs on plants
that already bear eggs and larvae; (3) Females might maintain the same mean fecundity
by laying more eggs per plant on plants lacking previously laid eggs or larvae; and/or
(4) Females might maintain the same mean fecundity by increasing the amount of time
spent searching for and ovipositing on A. reticulata. Each of these possible changes
would result in a decline in representational fitness, because of either a decline in
fecundity (1) or in offspring survivorship (2-4).7 Consequently, the fitness of non-
diapausers should decline as their frequency increases.
MAINTENANCE OF DIAPAUSE POLYMORPHISM
With the plausibility of frequency-dependence of representational fitness of non-
diapausers established, it is now possible to suggest how the diapause polymorphism
may be maintained in east Texas populations of Battus philenor. In this section I present
a simple genetic model of the evolution of diapause which shows that frequency-depen-
dence of the type seen in B. philenor can be sufficient to maintain a polymorphism.
The model assumes that whether an offspring of a first-brood female enters pupal
diapause or emerges to participate in the second brood is controlled by a single Mende-
lian locus with two alleles, A1 and A2. Individuals homozygous for A1 are non-diapaus-
ers, whereas individuals homozygous for A2 enter diapause. For simplicity, I treat in
detail the case in which heterozygotes also are non-diapausers (i.e., complete dominance
of allele A1), though it will be seen later that the properties of the model do not differ
in the case of complete recessiveness of A1, and hence, by inference, for cases of inter-
mediate dominance. I assume that genotype at the A locus affects only tendency to
diapause. In particular, I assume that the A-locus genotype does not affect mating
success, fecundity, or larval or pupal survivorship.
Let pi be the gene frequency of allele Ai, and let G1 be the genotype frequency of
An, G2 the genotype frequency of A12, and G3 the genotype frequency of A22 in overwin-
tering pupae. Assuming that random mating occurs among brood-1 individuals that
emerge from these pupae, the genotype frequencies among the offspring of brood-1
individuals are given by
G1 = pi2 (proportion of population that is homozygous, non-diapausing)
G2 = 2pip2 (proportion of population that is heterozygous, non-diapausing)
G, = P22 (proportion of population that is diapausing).
In particular, these are the genotype frequencies at the time of pupation when the
decision to diapause or not is presumably made. Consequently, a fraction p22 of the
individuals (those that are A22) enter diapause and the remainder, 1 p22, emerge to
form the second brood.
The gene frequencies, qi, among the non-diapausing portion of the population are
ql = (p,2 + P1P2)/(P12 + 21p2) = 1/(1 + P2) (see appendix footnote 8)
q2 = P1P2/(P 2 + 2pip2) = p/(l + P2).
Again, assuming random mating among non-diapausing individuals, the genotype fre-
quencies, Gi', among the offspring of second-brood individuals are
Florida Entomologist 69(1)
Gi' = q,2 = 1/(1 + p2)2
G2' = 2qiq2 = 2P2/(l + P2)2
G3' = q22 = 22/(1 + p2)2.
Next, let m be the mean number of eggs laid by brood-2 females and I be the mean
probability of survival of those eggs to the pupal stage. Then Wnd = Im is the represen-
tational fitness of non-diapausing females, as defined previously. Finally, let Wd be the
probability that a diapausing pupa will survive from the time brood-2 adults emerge
until their offspring pupate. The genotype frequencies among all overwintering pupae,
Gi", are then the weighted average of the frequencies among offspring of the first and
second broods, where the weightings are given by Wnd and Wd:
TG = (1 p2)Wnd/( + P2)2 (la)
TG2 = 2p2(l p22)Wnd/( + P2)2 (lb)
TG3 = p22(1 p22)Wd/(1 + p2)2 + PWd (ic)
and where T is the sum of the right-hand sides of the equations9, and is given by
T = (1 p22)Wnd + P22Wd (2).
Eqs. (1) are in fact the recursion equationsl0 for the system, since G3 + 1/2 G2 could be
substituted for p2. The recursion equation for the frequency of allele A2, obtained by
summing (Ic) and 1/2 of (Ib), is
Tp2 = P2( 2)Wnd + p22Wd (3).
In the previous sections, I have argued that the representational fitness of brood-2
females is frequency-dependent. In particular, Wnd is an increasing function of p22, the
proportion of diapausers. For simplicity, let this relationship be represented by Wnd =
kp22 + c, where k and c are constants. By contrast, there is little reason to suspect that
the survivorship of pupae is density-dependent and I therefore assume Wd is constant.11
I now show that if there is some value of P2 at which Wnd = Wd, then a stable
polymorphism will be maintained in the population. First note that if Wnd = Wd = W,
then T = W. This is shown by summing Eqs.(la)-(lc) and combining terms, recognizing
that the sum of the Gi is 1. When this is done, (3) reduces to
P2 = P2( P2) + P22 = P2,
which says that there is no change in gene frequency when Wnd = Wd. Because genotype
frequencies are functions of the gene frequencies, once gene frequencies reach equilib-
rium, so do genotype frequencies. The equilibrium gene frequency, p2, obtained by
Wd = kp22 + c,
P2 = V(Wd c)/k.
Next, I show that when Wnd < Wd, the frequency of the diapause allele, A2, will
increase, whereas if Wnd > Wd, that frequency will decrease. There will be an increase
in P2 (the frequency of allele A2 in one time period) if p2 (the frequency of allele A2 in
the next time period) P2 > 0, or, equivalently, if
Tp2 "- Tp2 > 0 (4).
Substituting (2) and (3) into (4) and simplifying yields Wd > Wnd. In similar fashion, it
can be shown that P2 decreases whenever Wd < Wnd.
In Fig.6 are portrayed the assumed relationships between Wd and the frequency of
diapausers, p22, and between Wnd and p22. The point at which these two curves intersect
represents an equilibrium, p22. That equilibrium is stable because for values of p22 less
than p22, Wnd < Wd, SO p22, and hence the frequency of A2, will increase toward the
equilibrium. By contrast, when p22 is greater than p22, Wnd > Wd and p22 will decrease
This analysis shows that the frequency-dependent effect of competition for oviposi-
tion sites on representational fitness can maintain a stable diapause polymorphism. The
only requirements are that (1) when the diapause allele is rare (i.e., when few individu-
Insect Behaviorial Ecology-'85 Rausher
0 P2 i
FREQUENCY OF DIAPAUSERS (p2)
Fig. 6 Assumed relationship between frequency of diapausers, P22, and fitness for
diapausers and non-diapausers. The broken line indicates the equilibrium frequency of
als diapause and competition for oviposition sites is intense), pupal survivorship during
the period when brood-2 adults are flying and their offspring are developing as larvae
is greater than the representational fitness of second-brood females (i.e., Wd > Wnd),
and (2) when the diapause allele is common (i.e., most individuals diapause and compe-
tition for oviposition sites is lax), pupal survivorship is less than the representational
fitness of second-brood females (i.e., Wd < Wnd). These two conditions ensure that the
two curves in Fig. 6 intersect and hence ensure the existence of a stable diapause
As presented, the analysis may seem restricted in its applicability because of the
assumption that A, is dominant to A2. If one performs a completely analogous analysis
under the assumption of A2 being dominant to A1, one still finds that requirements (1)
and (2) above guarantee the existence of a stable polymorphism.12 This and the previous
case represent the extremes in degree of dominance of one allele over the other, ignoring
cases of overdominance. Because in both cases requirements (1) and (2) above are
necessary and sufficient conditions for the existence of a stable diapause polymorphism,
this should also be true for intermediate degrees of dominance.
The existence of competition for oviposition sites among second-brood Battus
philenor females provides an explanation for the evolution and maintenance of a
diapause polymorphism in east Texas populations of this butterfly species. Because
oviposition sites are limited, an individual's expected genetic representation in the next
generation is inversely proportional to the number of females that compete for those
74 Florida Entomologist 69(1) March, 1986
A larva that pupates in mid- to late April in east Texas can be thought of as having
an option to either enter diapause and remain in that state until the following spring or
emerge two weeks later to participate in the second brood. Which option natural selec-
tion will favor depends crucially on what option other individuals in the population elect.
Consider first a situation in which all individuals in the population are genetically prog-
rammed to enter diapause. A mutant female that fails to enter diapause will then
emerge in early May and find an unexploited supply of Aristolochia serpentaria plants
on which to oviposit and on which its offspring may develop (provided, of course, a
mutant non-diapausing male is available for mating). Since there is no competition from
other females, the number of offspring of the mutant individual pupating successfully
should be high. In particular, as long as the expected number of offspring pupating is
is greater than the expected survivorship of diapausing pupae, the mutant type will
increase its genetic representation in the population as a whole, and the mutant will be
favored by selection.
As selection increases the frequency of the mutant type, however, the number of
females competing during the second brood will also increase, and consequently the
expected number of offspring per female reaching the pupal stage will decrease. This
decrease will continue until the expected number of offspring pupating per female
exactly equals the probability of survival of diapausing pupae. At this point, the genetic
contribution of a non-diapausing individual is exactly equal to that of a diapausing
individual and there will be no selective forces acting to increase or decrease the fre-
quency of non-diapausers. A similar argument holds for a mutant diapauser, which
would increase in a population of non-diapausers.
The model presented in the previous section is a formal representation of this
scenario. It assumes, of course, that diapause is controlled by a single Mendelian locus
with two alleles. Because nothing is currently known about the genetic control of
diapause in B. philenor, however, a polygenic mode of inheritance of diapause tendency
in B. philenor can not be ruled out; nor can the possibility be eliminated that the
decision to enter diapause is primarily environmentally controlled in B. philenor. In
many insects, a photoperiod threshold controls diapause. If individuals pupate when
daylength is below a certain threshold, a state of diapause is (or is not) entered. By
contrast, if pupation occurs with daylength above the threshold, diapause is not (is)
entered (Danilevsky et al. 1970, Tauber and Tauber 1976). Because there is at least a
2-3 week range of pupation dates for offspring of brood-1 B. philenor females in east
Texas (personal observation) during a time when daylength is rapidly increasing, it is
conceivable that the threshold daylength falls within that period. If so, individuals
pupating early will emerge to form a second brood while those pupating late will enter
Although the one-locus model presented above would not be relevant to a situation
in which diapause tendency were controlled either polygenically or environmentally, in
either case the basic verbal argument given above would still pertain. In fact, Slatkin
(1978) has shown that when the proportion of individuals in either of two phenotypic
classes is controlled polygenically, the genetic equilibrium is characterized by equilibra-
tion of the witnesses of the two phenotypic classes. Thus, Fig. 6 schematically portrays
the equilibrium for polygenic control of diapause, with the proviso that the x-axis is
taken to be the proportion of individuals in the population that enter diapause.
Environmental control of diapause is just a special case of polygenic control of
diapause, because the threshold daylength is likely to be genetically variable to some
degree and hence can presumably evolve (Hoy 1978). Increasing or decreasing the
threshold simply increases or decreases the proportion of the population that pupates
before daylength passes the threshold, and hence adjusts the proportions of individuals
that are in each phenotypic class (i.e., diapause or non-diapause). Slatkin's results are
Insect Behaviorial Ecology-'85 Rausher
then applicable and an equilibrium occurs when the fitness of non-diapausers (Wnd)
equals that of diapausers (Wd).
The explanation proposed here for the maintenance of a diapause polymorphism in
B. philenor is of course not the only one possible. In particular, it is conceivable that
this polymorphism is maintained by simple heterosis (Roughgarden 1979), by temporally
fluctuating selection pressures (Felsenstein 1976), or even represents an example of
"adaptive coin flipping" (Cooper and Kaplan 1982, Walker 1986). None of these pos-
sibilities, or even some combination of them, can be ruled out at this point. However,
demonstration of the existence of competition for oviposition sites and the reasonable
inference of frequency-dependence of the fitness of non-diapausers is positive evidence
arguing in favor of the explanation offered here. Closer scrutiny of the genetics and
ecology of diapause in Battus philenor will be needed to determine conclusively whether
that explanation is correct.
'Larvae fed a mixture of excised leaves of A. reticluata and A. serpentaria were
reared under natural temperature and photoperiod conditions on the porch of a cabin
in east Texas. Upon pupation, each larva was placed in its own container. If eclosion
occurred within three weeks of pupation, a butterfly was considered not to have entered
diapause. Individuals that failed to eclose within this period still had not closed within
six months, indicating that they were either in diapause or had died. These butterflies
emerged over the course of several months beginning in late December and early Jan-
uary in the laboratory. Only individuals that actually emerged were included in the
2Genetic variation for diapause tendency has been found in the papilionid Papilio
zelicaon. Simms (1983) argues that this variation is polygenic in nature, but his data
are also consistent with single-locus, Mendelian inheritance. Simms and Shapiro (1983)
show that environmental conditions can influence tendency to diapause in California
populations of Battus philenor, but they did not rule out the possibility that variation
in diapause tendency may also be explained partly by underlying genetic variation.
3In many insects, the rate at which females mature eggs, and hence the rate at which
eggs may be laid, declines as females age (Wigglesworth 1972, Price 1984). Because the
average age of females increases over the course of a brood (Odendaal, Lederhouse,
and Rausher, unpublished data), one might then expect to see a gradual decline in
average oviposition rate.
4In B. Philenor, the rate at which females alight on host plants is directly propor-
tional to the abundance of host plants in the habitat (Rausher 1983). If host plant
abundance decreases over time, then alighting rates will do likewise; and if the proba-
bility of oviposition is not influenced by alighting rate, then oviposition rate will also
decline. In east Texas, the number of host plants available to ovipositing females may
be expected to decline during brood 2 because they are consumed by dispersing larvae
that are the slowly growing offspring of brood-1 and the early-laid offspring of brood-2
5Because females tend to avoid ovipositing on plants that already bear eggs (Rausher
1979), the number of plants acceptable to females (i.e., that lack previously laid eggs)
is expected to decline over time. If oviposition probability, once a plant is alighted on,
remains constant, then oviposition rate will decline.
6During brood 1, there is no significant trend in the proportion of plants alighted on
that already bear eggs (Fig. 3a). This constancy is probably due to the variation in
host-plant leafing phenology, which causes the number of plants available to females to
increase steadily during the first brood (Rausher 1980, 1983). This increase in host-plant
numbers, along with the constancy in proportion of plants that bear eggs, is also respon-
sible for the steady increase in the number of host plants alighted on during the first
brood that do not bear eggs (Fig. 4a). Because there is an internally-regulated maximum
oviposition rate (Fig. la and Rausher 1983), while the number of host plants alighted
Florida Entomologist 69(1)
on rises steadily, the proportion of host plants alighted on that are accepted declines
steadily during the first brood (Fig. 5a; see also Rausher 1983).
7This conclusion is obvious for change (1). Since it is less obvious for the other
possible changes, I briefly provide here justification for why changes (2)-(4) would also
lower the representational fitness of non-diapausers. Rausher (1979) has shown that
offspring placed on plants that already harbor eggs or larvae have a lower survivorship
than eggs placed on unoccupied plants. Consequently, increasing the proportion of eggs
that are laid on occupied plants will decrease mean offspring survivorship, and hence
mean representational fitness.
Although we have no direct experimental evidence, it is also almost certain that
increasing the number of eggs laid per plant by a single female will also decrease
offspring survivorship. This conclusion is based on the results of several experiments
and observations. First, no single host plant is large enough to support the complete
development of one B. philenor larva. A larva, after feeding on all edible foliage on the
plant on which it hatches, must disperse to find other host plants (Rausher 1980, 1981,
Rausher and Papaj 1983b). The size of a larva when it disperses from its initial host
plant is directly proportional to the amount of edible foliage that was present on that
plant (Pilson and Rausher, unpublished). Presumably, the more larvae that are placed
on a plant, the less each obtains of the fixed amount of edible foliage, and the smaller
they are when they disperse from their first host. Because the probability of finding
another plant, and hence of surviving to the adult stage, decreases with decreasing size
at dispersal (Rausher 1979), an increase in the number of eggs a female lays on a host
plant will presumably decrease mean offspring survivorship, and hence decrease mean
Finally, an increase in the proportion of offspring placed on A. reticulata during the
second brood will also lower mean representational fitness. Because the leaves of A.
reticulata become sclerophyllous by the time of the second brood, whereas those of A.
serpentaria do not, the foliage of the latter host is much more suitable for larval develop-
ment than that of the former. Larvae that hatch on A. serpentaria are therefore much
larger when they disperse to find new hosts than larvae that hatch on A. reticulata,
resulting in greater overall survivorship for offspring placed on A. serpentaria (Rausher
1980, 1981). A shifting of eggs from A. serpentaria to A. reticulata in response to
increased competition for oviposition sites would therefore lead to a decrease in mean
offspring survival and in representational fitness.
"This identity may be derived as follows:
(pi2 + P1P2)/(P12 + 2pip2) = PI(P1 + P2)/PI(PI + 2p2)
= (P1 + P2)(p + P + P2)
and since p, + P2 = 1,
= 1/(1 + P2).
"T is a normalizing factor that, when divided into the right-hand side of Eqs. (1)
makes their sum equal to 1. By multiplying each side of these equations by T, one
obtains the form shown in the text.
'0A set.of recursion equations provides the mathematical rule that transforms gene
or genotype frequencies in one time period into the frequencies in the next time period.
"The inferences derived from the model remain valid even if density-dependence
exists, as long as the slope of the relationship between pupal survival and P22 is less
than k, i.e, as long as the curves relating gene frequency to fitness for diapausers and
non-diapausers intersect and to the right of the intersection (high values of p2) the curve
for non-diapausers is above that for diapausers.
"In this case the relationship between gene frequency and representational fitness
is assumed to be
Wnd = k(1 pi2) + c
and the equilibrium gene frequency is
P2 = 1 V(k + c Wd)/k.
Insect Behaviorial Ecology-'85 Rausher
I wish to thank Jim Lloyd for organizing the symposium at which this paper was
presented. I also wish to thank Frank Slansky, Thomas Walker, and an anonymous
reviewer for suggested improvements to the manuscript. Anne Lacey contributed in-
valuable moral support throughout the writing of this manuscript. This work was sup-
ported in part by NSF grant BSR-8406870.
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CALLING BEHAVIOR: CAN IT BE USED TO
IDENTIFY MIGRATORY SPECIES OF MOTHS?
JEREMY N. MCNEIL*
Department de Biologie
Ste. Foy, P.Q., GlK 7P4
There is considerable debate concerning the fate of populations of noctuid moths,
founded during the summer, by migrants moving considerable distances northward of
areas where permanent populations persist. This paper compares the female calling
behavior of moth species that maintain permanent populations in Canada with those
that are known immigrants. Residents initiate calling soon after emergence, regardless
of climatic conditions, while the mean age of calling for immigrant species is variable,
even under favorable conditions. Furthermore the onset of calling by immigrants is
*Jeremy McNeil is a full professor in the Biology Department at Laval University. His main interests are the
ecological and behavioral aspects of pheromone mediated systems in insects, especially Lepidoptera. Current address:
Department de biologie, Universite Laval, Ste. Foy, P.Q., G1K 7P4, Canada. (Contribution 470, Departement de
Insect Behaviorial Ecology-'85 McNeil 79
strongly influenced by prevailing temperature and photoperiodic conditions during adult
life. These differences may only be of importance for the northward spring dispersal,
but as fall conditions induce a delay in the onset of calling of known immigrants it is
proposed that this could potentially permit a southerly, upper air migration to suitable
A number of noctuid moths of major economic importance in North America (see
Rabb and Kennedy 1979) are known to be highly mobile and, during the summer
months, expand their geographic distribution into northerly areas where they are un-
able to establish permanent populations. These temporary populations have often been
considered as dead end ones that are killed off at the onset of adverse weather conditions
in the fall. Rabb and Stinner (1978) referred to this type of movement as the "Pied
Piper" phenomenon, suggesting that man's agricultural practices temporarily increase
suitable and available habitats and that "this [the Pied Piper effect] seems to "pervert"
the survival strategies of the species involved. . .". However, Walker (1980) proposed
that if there is a genetic basis for this repeated seasonal dispersal, the demise of indi-
viduals exploiting these temporally available habitats would create an evolutionary di-
lemma as such suicidal dispersal would be strongly selected against. Stinner et al. (1983)
provided a counter argument on this point, but agreed that the testable predictions
proposed by Walker (1980) would help elucidate whether or not noctuids have a return
There is considerable concrete evidence that some butterfly species have unidirec-
tional spring and fall migrations, generally occurring within the boundary layer (Walker
1980; Baker 1984 and references therein). On the other hand there is little data for night
flying moths, one obvious reason being the difficulty with which direct observations can
be made. Two well documented cases for noctuids do exist: the bogong moth in Australia
(Common 1954) and the army cutworm in North America (Pruess 1967; Kendall 1981)
migrate, as adults in reproductive diapause, to the mountains for the summer before
returning in the fall to the plains where they reproduce. Dingle (1982) pointed out that
migration associated with adult reproductive diapause is quite common, and the derived
advantages of such a system are evident for Lepidoptera that migrate to aestivation
(the army cutworm) or overwintering sites (the monarch) and return before the onset
of reproduction. However, temperate species that expand their range into temporarily
available habitats during the summer months, if capable of a return fall migration,
would benefit from pre-reproductive strategies permitting the optimal utilisation of
both summer and winter resources. Based on data pertaining to the calling behavior of
several temperate species of Lepidoptera, I propose that certain insects undertaking
long distance, upper air, northward migrations (i) possess pre-reproductive traits that
differ considerably from resident species and (ii) that these traits, which fit within the
context of the diapause syndrome (Tauber et al. 1984), would facilitate a return south-
ward migration to suitable overwintering sites in the fall.
The true armyworm, Pseudaletia unipuncta is a noctuid that is found annually
throughout much of eastern Canada but does not establish permanent populations due
to the inability to overwinter in these areas (Ayre 1985; Fields and McNeil 1984). The
closely synchronized appearance of adults over a wide area, having very different local
climatic conditions, strongly supports the hypothesis that moths immigrate on prevail-
Florida Entomologist 69(1)
ing weather fronts, as suggested for other Lepidoptera (Arthur and Bauer 1981; Domino
et al. 1983). Studies investigating the nocturnal calling behavior of true armyworm
virgin females has shown that even under favorable summer conditions (25C, 16L:8D)
individuals initiate calling for the first time from 2 to 12 days following emergence; the
mean age for the onset of calling varying from 4 to 6 days (Turgeon and McNeil 1982;
Delisle and McNeil 1986). The mean calling age of P. unipuncta is also significantly
influenced by ambient temperature conditions during adult life (Turgeon and McNeil
1983); the lower the temperature, the older females are when they start calling, e.g. at
100C the mean age of calling is 17.9 days. Furthermore there is a photoperiodic effect,
with females at 25C calling significantly later under 12L:12D (X=7.9 days) than at
16L:8D (X=6.0 days) (Delisle and McNeil 1986). The combined effect of 10"C, 12L:12D,
during adult life resulted in a mean calling age of approximately 21 days, and when
females were transferred to 250C, 16L:8D, at 5, 10, or 15 days following emergence,
calling was initiated within six days (Delisle and McNeil, unpublished). The observed
delays in the onset of calling fit the "oogenesis-flight syndrome" (Johnson 1969), being
initiated by short days and low temperatures, conditions responsible for the initiation
of diapause of many temperate species (Beck 1980). I believe the responses observed
in the true armyworm reflect one extreme within the spectrum of adult reproductive
diapause associated with the migration of Lepidoptera as (i) ovarian development does
occur, albeit slowly, under short day/low temperature conditions and, (ii) females in-
itiate calling rapidly when transferred to long day/high temperature conditions. As
previously stated this strategy would benefit species that reproduce in both the summer
and winter limits of the distribution. This contrasts with the other extreme reported in
other noctuids, such as the army cutworm and the bogong moth (Common 1954; Pruess
1967), where reproductive diapause persists for several months. In these cases, little
or no reproductive activity occurs before the adults that immigrated to the mountains
initiate their return migration.
A markedly different situation is seen with the Bertha armyworm, Mamestra con-
figurata, a species that does overwinter in Canada. For this moth temperature and
photoperiodic conditions have very little effect on the age at which calling starts. At
25C, 16L:8D, all females initiated calling by the fourth night following emergence, with
a mean age of first calling of 2.2 days (Howlander 1985; J. Haley, unpublished data),
while at 10C, 16L:8D, it is 5.2 days (Howlander 1985). In addition, 100% of all females
held at 250, 12L:12D, were calling by the second night following emergence (Howlander
While the data base relative to calling behavior under different abiotic conditions is
less substantial, there is evidence that differences similar to those observed between
the true armyworm and the Bertha armyworm may exist for other resident and immig-
rant noctuids. Greater than 83% of virgin females of the potato stem borer, Hydraecia
micacea, an introduced European species that is now an established resident in Canada,
called the first night following emergence at 23C, 14L:10D, in the laboratory, while
100% did so under field conditions (West et al. 1984). On the other hand Swier et al.
(1976) reported that at 24-27C under 16L:8D, the black cutworm, Agrotis ipsilon, a
suspected spring immigrant (Domino et al. 1983; Kaster and Showers 1982), had an
average precopulatory period of 4.4 days, and that it took at least three nights following
emergence for more that 33% of the virgin females to initiate calling under a 18L:6D
photoperiod at 24C (Swier et al. 1977). A high incidence of unmated females in fall light
trap catches, together with low male catches in pheromone traps, was interpreted as
evidence of a reproductive diapause that might facilitate a return fall migration of the
black cutworm (Kaster and Showers 1982). Similar results have been collected since
1979 for the true armyworm in our Quebec trapping program (McNeil, unpublished),
and the delay in the onset of reproductive activity has been proposed as potentially
Insect Behaviorial Ecology-'85 McNeil 81
facilitating a return migration of this species to suitable overwintering sites (McNeil
1986). Consequently, I suggest that the black cutworm responds to fall conditions in
the same way as the true armyworm, with only a short delay in the onset of reproduc-
tion. Examination of the calling behavior of A. ipsilon under a range of different climatic
conditions would test this hypothesis.
In an effort to determine whether the proposed differences between resident and
migrant species have any wider application within the Lepidoptera my students and I
have recently started comparative work on the calling behavior of pyralids, because
members of this Family, like noctuids, are capable of upper air migration (Drake 1985).
The sunflower moth, Homoeosoma electellum, which migrates annually into Canada
(Arthur and Bauer 1981), was chosen as the migrant species as it has a number of
biological attributes that differ considerably from the true armyworm. The sunflower
moth calls during the photophase (Arthur 1978) rather than the scotophase, even though
for most other activities this species is nocturnal, and has the ability to enter diapause
as a last instar larva (Chippendale and Kikukawa 1983; Kikukawa and Chippendale
1983; Teetes et al. 1969). However, in Canada non-diapausing larvae leave the host in
August and give rise to adults in September. At this time climatic conditions do not
permit the completion of another generation, making it highly unlikely that H. electel-
lum successfully overwinters this far north (Arthur 1978). If this is the case then these
adults, like those of the true armyworm, would have to migrate southward if their
offspring are to survive. The resident species chosen for this study was the European
corn borer, Ostrinia nubilalis, which like the sunflower moth overwinters as a last
instar larva. While studies have only recently been initiated, the data obtained to date
support the hypothesis that the pre-reproductive behavior of migrant and resident
temperate Lepidoptera vary considerably. Females of both univoltine and bivoltine
races of the European corn borer call within two days of emergence at 25C, 16L:8D
(L. Royer and McNeil, unpublished), while under the same conditions sunflower moth
females have a mean age of calling of 9.7 days (McNeil and Delisle, unpublished). The
longevity of our H. electellum adults, established from immigrants collected in Saska-
toon, while similar to that of the Missouri strain (Kikuwawa and Chippendale 1983),
was considerably longer than the 8.5 days reported for adults from Texas (Randolph et
al. 1972). Temperature and photoperiodic conditions were similar in the laboratory
hearings, although other aspects such as relative humidity, available food, and adult
densities could explain the observed differences in adult longevity. However, an alter-
nate explanation that merits further investigation within the context of the migration
of this species is the possible existence of polymorphism in the population, where life
statistics of migrants exploiting temporarily available habitats differ from those that
remain in the area of permanent occupation.
The evidence presented above strongly supports the hypothesis that pre-reproduc-
tive strategies of resident and migrant moth species differ markedly. Indigenous species
mate soon after emergence, which would permit rapid exploitation of the resources
available in the immediate habitat. On the other hand, species that have significant
northward expansions in their summer distribution show considerable variability in the
time required for the onset of reproduction, even under optimal summer conditions. At
least in the case of the true armyworm, and most probably in the case of the black
cutworm, fall conditions induce a. further delay in reproductive activity. This could
provide the time necessary for a southerly migration to favorable overwintering sites
if dispersal occurred in upper air masses, where it has been shown that moths may be
carried for at least 90 km in 24 h (Rose et al. 1985). Furthermore, the rapidity with
which sexual activity is resumed when females are transferred to favorable conditions
82 Florida Entomologist 69(1) March, 1986
would permit successful emigrants to maximize their reproductive potential upon arrival
in acceptable overwintering habitats. However, whether or not seasonal variability in
pre-reproductive behavior is uniquely associated with emigration from sites where per-
manent populations occur or, as suggested by McNeil (1986), is a life history trait
permitting a return migration remains to be elucidated. There is one encouraging point
for those of us who believe that the answer to Walker's question, "are butterflies better
than moths?" (Walker 1980), is no. The oriental armyworm, P. separate, the only noc-
tuid species where clear evidence for both north and south migration has been obtained
by means of a massive adult mark-recapture program (Li et at. 1964, reference from
Baker 1978), has at least a 3 day delay in the onset of calling after emergence at 250C,
16L:8D (Hirai 1984), similar to P. unipuncta.
I would like to thank Jim Lloyd for the kind invitation to participate in the 1985
Behavioral Ecology Symposium. Thanks are also due to M. Cusson, J. Delisle, S.
Fitzpatrick and J. Richardson for their comments on an earlier version of this manu-
script, as well as to T. J. Walker for valuable editorial suggestions.
ARTHUR, A. P. 1978. The occurrence, life history, courtship, and mating behaviour
of the sunflower moth, Homoeosoma electellum (Lepidoptera: Phycitidae) in the
Canadian prairie provinces. Canadian Ent. 110: 913-16.
ARTHUR, A. P., AND D. J. BAUER. 1981. Evidence of the northerly dispersal of the
sunflower moth by warm winds. Environ. Ent. 10: 528-33.
AYRE, G. L. 1985. Cold tolerance of Pseudeletia unipuncta and Peridroma saucia
(Lepidoptera: Noctuidae). Canadian Ent. 117: 1055-60.
BAKER, R. R. 1978. The evolutionary ecology of animal migration. Holmes and Meier,
1984. The dilemma: When and how to go or stay. Pages 279-296 In R. I.
Vane-Wright and P. R. Ackery, Eds. The biology of butterflies. Academic Press,
BECK, S. D. 1980. Insect photoperiodism. 2nd Edition. Academic Press, New York.
CHIPPENDALE, G. M., AND S. KIKUKAWA. 1983. The effect of daylength and tem-
perature on the larval diapause of the sunflower moth, Homoeosoma electellum.
J. Insect Physiol. 29: 643-49
COMMON, I. F. B. 1954. A study of the ecology of the adult bogong moth, Agrostis
infusa (Boesd.) (Lepidoptera: Noctuidae), with special reference to its behaviour
during migration and aestivation. Australian J. Zool. 2: 223-63.
DELISLE, J., AND J. N. MCNEIL. 1986. The effect of photoperiod on the calling
behaviour of virgin females of the true armyworm, Pseudalatia unipuncta. J.
Insect Physiol. (In press).
DINGLE, H. 1982. Function of migration in the seasonal synchronization of insects.
Ent. Expt. Appl. 31: 36-48.
DOMINO, R. P., W. B. SHOWERS, S. ELWYNN TAYLOR, AND R. H. SHAW. 1983.
Spring weather pattern associated with suspected black cutworm moth
(Lepidoptera: Noctuidae) introduction to Iowa. Environ. Ent. 12: 1863-72.
DRAKE, V. A. 1985. Radar observations of moths migrating in a nocturnal low-level
jet. Ecol. Ent. 10: 259-65.
FIELDS, P. G., AND J. N. MCNEIL. 1984. The overwintering potential of the true
Insect Behaviorial Ecology-'85 McNeil 83
armyworm, Pseudaletia unipuncta (Lepidoptera: Noctuidae), populations in
Quebec. Canadian Ent. 116: 1647-52.
HIRAI, K. 1984. Migration of Pseudaletia separate Walker (Lepidoptera: Noctuidae).
Considerations of factors affecting time of taking-off and flight period. Ann. Ent.
Zool. 19: 422-29.
HOWLANDER, M. M. A. 1985. The biology of calling behaviour in the Bertha ar-
myworm, Mamestra configurata Walker (Lepidoptera: Noctuidae). Ph.D.
Thesis, University of Manitoba.
JOHNSON, C. G. 1969. Migration and disperal of insects by flight. Metheun, London.
KASTER, L. V., AND W. B. SHOWERS. 1982. Evidence of spring immigration and
autumn reproductive diapause of the adult black cutworm in Iowa. Environ. Ent.
KENDALL, D. M. 1981. Bionomics of Euxoa auxiliaris Grote (Lepidoptera: Noc-
tuidae) in the Rocky Mountains and comparison with two resident species of
alpine moths. M. Sc. Thesis, University of Colorado.
KIKUKAWA, S., AND G. M. CHIPPENDALE. 1983. Seasonal adaptations of different
geographic populations of the sunflower moth, Homoeosoma electellum. J. Insect
Physiol. 30: 451-55.
MCNEIL, J. N. 1986. The true armyworm Pseudaletia unipuncta (Haw.) (Lepidopt-
era: Noctuidae). A possible migrant species. In D. R. MacKenzie, C. S. Barfield,
G. G. Kennedy and R. D. Berger, Eds. The movement and dispersal of agricul-
turally important biotic agents. Claitor's Publishing Division, Baton Rouge (in
PRUESS, K. P. 1967. Migration of the army cutworm, Chorizogrotis auxiliaries
(Lepidoptera: Noctuidae). I. Evidence of a migration. Ann. Ent. Soc. America
RABB, R. L., AND G. G. KENNEDY, Eds. 1979. Movement of highly mobile insects:
Concepts and methodology in research. North Carolina State University,
RABB, R. L., AND R. E. STINNER. 1978. The role of insect dispersal in population
processes. Pages 3-16 In C. R. Vaughn, W. Wolf and W. Klassen, Eds. Radar,
insect population ecology and pest management. NASA Conf. Publ. No. 2070,
NASA Wallops Flight Center, Wallops Island, VA.
RANDOLPH, N. M., G. L. TEETES, AND M. C. BAXTER. 1972. Life cycle of the
sunflower moth under laboratory and field conditions. Ann. Ent. Soc. America
ROSE, D. J. W., W. W. PAGE, C. F. DEWHURST, J. R. RILEY, D. R. REYNOLDS, D.
E. PEDGLEY, AND M. R. TUCKER. 1985. Downwind migration of the African
armyworm moth, Spodoptera exempta, studied by mark-and-capture and by
radar. Ecol. Ent. 10: 299-313.
STINNER, R. E., C. S. BARFIELD, J. L. STIMAC, AND L. DOHSE. 1983. Dispersal
and movement of insect pests. Annu. Rev. Ent. 28: 319-35.
SWIER, R. W. RINGS, AND G. J. MUSICK. 1976. Reproductive behavior of the black
cutworm, Agrotis ipsilon. Ann. Ent. Soc. America 69: 546-50.
1977. Age-related calling behavior of the black cutworm, Agrotis ipsilon. Ann.
Ent. Soc. America 70: 919-24.
TAUBER, M. J., C. A. TAUBER, AND S. MASAKI. 1984. Adaptations to hazardous
seasonal conditions: Dormancy, migration and polyphenism. Pages 149-83. In C.
B. Huffaker and R. L. Rabb, Eds. Ecological entomology. John Wiley and Sons,
TEETES, G. L., P. L. ADKISSON, AND N. M. RANDOLPH. 1969. Photoperiod and
temperature as factors controlling the diapause of the sunflower moth,
84 Florida Entomologist 69(1) March, 1986
Homoeosoma electellum. J. Insect Physiol. 15: 755-61.
TURGEON, J. J., AND J. N. MCNEIL. 1982. Calling behaviour of the armyworm,
Pseudaletia unipuncta. Ent. Expt. Appl. 31: 402-8.
1983. Modifications in the calling behaviour of Pseudaletia unipuncta (Haw.)
(Lepidoptera: Noctuidae), induced by temperature conditions during pupal and
adult development. Canadian Ent. 115: 1015-22.
WALKER, T. J. 1980. Migrating Lepidoptera: Are butterflies better than moths? Flor-
ida Ent. 63: 79-98.
WEST, R. J., P. E. A. TEAL, J. E. LAING, AND G. M. GRANT. 1984. Calling behavior
of the potato stem borer, Hydraecia micacea Esper (Lepidoptera: Noctuidae) in
the laboratory and the field. Environ. Ent. 13: 1399-1404.
PREZYGOTIC MALE REPRODUCTIVE EFFORT IN INSECTS:
WHY DO MALES PROVIDE MORE THAN SPERM?
JAMES S. QUINN* AND SCOTT K. SAKALUK*
Males' incentives for providing benefits to females and/or their offspring are ambigu-
ous during the period prior to zygote formation. The benefits may function to increase
the number of available eggs fertilized by a male and/or enhance the production and
survival of his offspring. In some cases, male prezygotic investment may be an adapta-
tion to secure fertilizations despite the fact that it incidentally benefits the female or
her offspring. More often, the benefits to offspring production and survival are not
simply incidental and probably account, in part, for the magnitude of the male invest-
ment. Regardless of the adaptive significance of male provided benefits, they typically
reduce the females' costs of producing surviving offspring while raising the males' costs.
The extent to which provisioning of benefits increases males' costs and decreases
females' costs will affect the degree to which females limit male reproduction (or vice
versa). If male-provided benefits (prezygotic or otherwise) are more costly than female
costs of offspring production, reproductively-ready males will act as resources limiting
female reproduction. From an evolutionary perspective it is important to consider the
effect of male-provided benefits. The primary function of the investment (e.g., to
maximize sperm transfer) is irrelevant in terms of the degree to which one sex limits
the other's reproduction.
"Women and men move back and forth in between effect and
cause. Just beyond the range of normal sight, this glittering
joker was dancing in the dragon's jaws"
During the past two decades there has been considerable interest in Darwin's (1871)
*James S. Quinn is a Ph.D. student at the University of Oklahoma with research interest in behavioral ecology
and is currently studying the causes and consequences of sexual size iln...rl.h.r,, ,, ,. .rrn. -h .
compared with Caspian Terns (Sterna caspia). Scott K. Sakaluk is a !...i -l.. r..i irl .. ir .- r .. i, .ri ..r ..
with research interests in insect reproductive behavior and especially the evolution of cricket mating systems.
Current addresses: Quinn, Department of Zoology, University of Oklahoma, Norman, OK 73019; Sakaluk, Depart-
ment of Entomology, University of Arizona, Tueson, AZ 85721.
Insect Behaviorial Ecology-'85 Quinn & Sakaluk 85
theory of sexual selection. In various theoretical discussions, authors have attempted
to identify the key variables affecting the operation and intensity of sexual selection
(Trivers 1972, Emlen and Oring 1977, Wade and Arnold 1980, Gwynne 1984a, Thornhill
in press. At the same time, other studies have investigated the mechanisms through
which sexual selection is mediated, specifically, intrasexual competition and mate choice
(for a review of these processes in insects see Thornhill and Alcock 1983). These two
approaches ask very different questions-what causes sexual selection? and what is the
function of adaptations that arise through selection processes? In this paper we will
adopt Williams' (1966) use of "function" to mean that the characteristic being considered
was fashioned by natural selection for the goal attributed to it. When such a relationship
is not intended we will use the term "effect" to imply that certain consequences of the
characteristic may not necessarily be the goal fashioned by natural selection (see Wil-
Relative parental contribution by the sexes has long been considered a key variable
influencing the operation of sexual selection. Bateman (1948) proposed that the unequal
energetic expenses of gamete production led to the typical pattern of courtship in ani-
mals. He found that in Drosophila melanogaster, female reproduction was limited by
the number of eggs laid while male reproduction was limited by the number of females
inseminated. Thus, females could be viewed as a limiting resource for which males were
expected to compete. Females are selected to be more discriminating in their choice of
mates because they have more to lose (their initial gametic investment) from mating
with a genetically incompatible or otherwise unsuitable mate.
Trivers (1972) provided a more general hypothesis to explain interspecific differences
in the operation of sexual selection, proposing that the amount of parental investment
by the sexes was the key variable influencing the operation of sexual selection. He
defined parental investment as "any investment by the parent in an individual offspring
that increases the offspring's chance of surviving (and hence reproductive success) at
the cost of the parent's ability to invest in other offspring." Trivers concluded that
parental investment, measured as a cost to other parenting efforts, ultimately explained
sexual differences in such life history variables as age of first breeding, differential
mortality, and adult sex ratio, as well as the differences in sex roles during mating.
Insect females typically provide almost all parental investment in their production
of eggs. However, in some species parental investment is shared. An example of paren-
tal investment by a male insect comes from the work of R. L. Smith (1979a, 1979b,
1980) in his studies of the giant waterbug Abedus herberti, and other Belostomatidae.
These males provide ovipostion sites (on their backs) for the eggs of their mates and
provide care for those eggs. Paternal care involves aeration of eggs and brood pumping,
a behavior that expedites the escape of young from the egg. This paternal care involves
a cost to the male's ability to invest in other offspring because back space is limited.
The extra drag associated with swimming while encumbered, as well as brooding be-
haviors, may increase risks of predation and decrease success at capturing prey (Smith
1979b, 1980). This increased time and risk associated with paternal care excludes encum-
bered males from mating and apparently affects the operational sex ratio (Emlen and
Oring 1977), as well as the courtship roles.
In this paper, we consider costly investments by male insects before they fertilize
the eggs of females. We investigate the adaptive significance of such prezygotic invest-
ments, and examine the impact of these benefits on the operation of sexual selection.
We consider the contention that prezygotic male investment is best considered mating
effort (Low 1978) and not parental investment. We conclude that from an evolutionary
perspective, the effects of the investment, and not the primary function, are the critical
Florida Entomologist 69(1)
CATEGORIES OF MALE PREZYGOTIC INVESTMENT
Males often minimize their investment in offspring. This is because males produce
many energetically inexpensive sperm compared with few expensive eggs produced by
females (Bateman 1948). Male reproduction is typically limtied by the number of females
inseminated, a potentially great number. By providing parental effort rather than seek-
ing other copulations, males potentially experience greater losses of other reproductive
opportunities than would females who make additional investments in offspring. Addi-
tionally, males may not be certain of the paternity of their mate's offspring and therefore
risk investing in the offspring of other males (Thornhill and Alcock 1983). Despite the
substantial costs, males of many insect species often provide benefits to females before
the females's eggs are fertilized. Such prezygotic investments include:
1. nuptial prey items consumed by the female (Downes 1970, Thornhill 1976a, 1980,
1983 and references).
2. spermatophores and/or accessory gland secretions
a. ingested orally by the female (Boldyrev 1927, Alexander and Otte 1967a,
Mullins and Keil 1980, Sakaluk and Cade 1980, Gwynne 1981, 1983, 1984b,
Bowen et al. 1984, Sakaluk 1984, 1985)
b. absorbed in the female's reproductive tract (Friedel and Gillott 1977, Boggs
and Gilbert 1979, Sivinski 1980a, Boggs 1981, Greenfield 1982, Marshall 1982,
Rutowski 1982, Schal and Bell 1982, Markow and Ankney 1984, Pivnick and
3. secretions from glands other than the accessory glands (Fulton 1915, Mays 1971,
Walker 1978, Bell 1980a,b, Bidochka and Snedden 1985)
4. portions of the male's body (Alexander and Otte 1967b, Dodson et al. 1983,
5. protection of the female from predators (Sivinski 1980b, 1983)
6. reduction of harassment of the female by other conspecific males (Waage 1979,
1983, Borgia 1981, Wilcox 1984)
7. assistance in the collection of suitable oviposition substrate such as dung or carr-
ion (Klemperer 1983, Tyndale-Biscoe 1984, Wilson and Fudge 1984)
8. provisioning of a burrow in which the female can rear her offspring (Walker 1980,
Trivers specifically excluded effort spent finding or subduing a mate from his concept
of parental investment, except in those cases where such effort affects the survival
chances of the offspring. For example, defense of a territory that benefits offspring
survival, providing nuptial gifts, and other behaviors providing "incidental" benefits to
offspring were included as parental investment. Low (1978) proposed that reproductive
effort be divided into two components, mating effort and parental effort. The division
and attendant definitions have resulted in a roundabout redefinition of parental invest-
ment that stresses the primary function of the investment rather than any effects it
might have on offspring survival or production costs.'
Alexander and Borgia (1979) and Gwynne (1984a) continued to stress the primary
function of the components of reproductive effort (Low 1978). The former argued that
because males have little control over the fate of their gametes or their mates' uses of
prezygotic paternal contributions, such investments would be best considered mating
effort. Gwynne (1984a) similarly identified prezygotic reproductive effort by males as
mating effort2 and suggested a further subdivision to distinguish effort influencing the
operation of sexual selection. His "non-promiscuous mating effort" refers to effort pro-
viding benefits to offspring or mates at a cost of lost reproductive opportunities, while
"promiscuous mating effort" provides no such benefits (Gwynne 1984a). Non-promiscu-
ous mating effort, as defined, is similar to parental investment in its effect on sexual
Insect Behaviorial Ecology-'85 Quinn & Sakaluk 87
selection. However, in species where males offer no more than sperm to females, sub-
stantial male investment in courtship, guarding, or fighting (promiscuous mating effort)
also can influence the operation of sexual selection and lead to male mate choice or even
sex role reversal in courtship behavior (Hatziolos and Caldwell 1983, Johnson and Hub-
bell 1984 and references). Furthermore, as mentioned earlier, the classification of such
effort as a type of mating effort is based on the primary function of the effort or the
"intention" of the male.
These different treatments of prezygotic male reproductive effort can result in the
identification of an effort that benefits offspring as either a parental investment (Trivers
1972, Thornhill 1976b, Boggs and Gilbert 1979, Morris 1979, Mullins and Keil 1980, Zeh
and Smith 1985) or a mating effort (Alexander and Borgia 1979, Gwynne 1984a, Thor-
nhill in press). The basis for this difference is whether male-provided benefits are
viewed according to the effect on offspring production or according to the primary
function of the investment.
ADAPTIVE SIGNIFICANCE: WHY Do MALES PROVIDE MORE THAN SPERM?
The selective advantages to males providing benefits to mates and offspring include
two main categories. First, such investments may increase the number of eggs fertilized
by the male's sperm by: a) increased sperm transfer to the female; b) increased utiliza-
tion of that male's sperm by the female; and/or c) increased number of females insemi-
nated. Second, male-provided benefits may increase the production and survival of
offspring. Male-derived selective advantages in the second category would be devalued
if some or all of the benefited offspring were fathered by other males".
The evolution of beneficial prezygotic investments by males is more likely in species
in which males can provide useful services (e.g., protection from predators) or collect
and/or defend resources that potentially limit reproduction. In species where reproduc-
tion is limited by protein or nutrients that are contained within the sperm or sperm
transfer device, females are expected to digest any materials as long as this does not
result in costly infertility of eggs. Thus, particularly in species engaging in multiple
mating, females may make use of sperm as a nutritional resource.
Many of the examples of male prezygotic investment provide both increased fertili-
zations and increased survival or production of offspring. For example, mate guarding
by male damselflies and dragonflies increases the male's likelihood of fertilizing eggs,
diminishes predation on the mate (and hence her offspring) and increases oviposition by
the female by reducing harassment by other males (Waage 1979, 1983). Detailed exam-
ples from crickets (Orthoptera: Gryllidae) and katydids (Orthoptera: Tettigoniidae) will
help illustrate the functions and effects of male prezygotic contributions.
First consider the adaptive significance of the spermatophylax produced by male
decorated crickets. Gryllodes supplicans (Sakaluk 1984, 1985). In crickets, copulation
ends when a male transfers the sperm-containing vessel (spermatophore) to the female.
The spermatophore of most cricket species consists of a small sperm-containing ampulla
that remains outside the female's body after mating. This ampulla is drained as the
sperm enter the female genital tract. Later the female often eats the evacuated sper-
matophore (Alexander and Otte 1967a, Loher and Rence 1978, Sakaluk and Cade 1980,
1983), but premature removal is prevented by the male who remains with and anten-
nates his mate (Loher and Rence 1978). Interestingly, G. supplicans males produce a
bipartite spermatophore consisting of the ampulla plus a spermatophylax, a larger
gelatinous portion devoid of sperm (Alexander and Otte 1967a, Sakaluk 1984, Sakaluk
and O'Day 1984). Immediately after mating, the female removes the easily detached
spermatophylax from the ampulla with her mouthparts and begins to feed on it. The
time required for a female to consume this nuptial 'meal' completely, increases with
Florida Entomologist 69(1)
spermatophore weight (Sakaluk 1985). Within several minutes of eating the sper-
matophylax, the female removes and eats the sperm ampulla and the remaining con-
tents. Thus a male that provides a small spermatophylax will have his ampulla removed
sooner than a male providing a larger one. Because the ampulla must be attached for
a minimum of about 50 min to be emptied completely of sperm (Sakaluk 1984), males
providing under-sized spermatophylaxes will not transfer a full complement of sperm.
However, the average time at which females removed sperm ampullae was 52 min,
which matches the time required for complete sperm transfer (Sakaluk 1984). Therfore
males, on the average, provide females with a nuptial meal no larger than that required
to prevent the premature removal of the ampulla. The ease with which the sper-
matophylax breaks off is important to fulfilling the function of preventing premature
removal of the ampulla. Furthermore, male G. supplicans are definitely less intense in
their post-copulatory interaction with the female than other crickets, which are known
to rely on direct contact and antennation for prevention of premature ampulla-removal
by females (Loher and Rence 1978). This suggests that the spermatophylax "replaces"
contact guarding and that the function of the bipartite spermatophore is to maximize
sperm transfer (Alexander and Otte 1967a).
The largest possible sperm transfer is expected to be particularly important in
species where sperm from a number of males are mixed in the spermatheca and fertili-
zation success is essentially by lottery (Parker 1970, Sakaluk in press). It appears that
the function of the spermatophylax, or more specifically the bipartite nature of the
spermatophore, is to ensure the maximum transfer of a male's sperm. There are other
possible beneficial effects of this nuptial offering, such as increasing production and
survival of offspring or inducing a female refractory period (i.e., a period when the
female refrains from mating). However, the bipartite spermatophore of G. supplicans
was not likely designed by natural selection to provide these other possible benefits.
There may be additons to the spermatophore that serve those other functions, such as
substances included in the spermatophore that promote a female refractory period or
that provide nutrition for the young. It appears likely that the evolution of the sper-
matophylax itself did not require males to reap these extra benefits. Thus, some of
these benefits may be incidental to the evolution of the spermatophylax.
The size of a spermatophylax represents a continuous variable. Costs of producing
a spermatophore are not trivial; in G. supplicans, the spermatophore can assume up to
about 6% of the male's body weight and males require 3.3-0.1 h for spermatophore
replenishment before they can remate (Sakaluk 1985). This may be a considerable cost
if mating opportunities are lost during this period of replenishment. In other crickets,
males transfer small unipartite spermatophores consisting of a sperm ampulla alone and
remate within as little as 15 min (Alexander and Otte 1967a). The costs can be viewed
as time or energy, but ultimately as lost reproductive opportunity. These costs are
weighed against the benefits-increased likelihood of fertilizing available eggs and in-
creased number of surviving offspring. In G. supplicans, secondary benefits to in-
creased spermatophylax size, such as potentially increased production of surviving
offspring, may be small. Nevertheless, the determination of the optimal spermatophylax
size by natural selection involved the balancing of all benefits agianst all costs.
Research by D. T. Gwynne and colleagues (Gwynne 1984b, Gwynne et al. 1984,
Bowen et al. 1984) on the katydid Requena verticalis serves to illustrate the impact of
secondary benefits on optimal investment in a continuous character. The bipartite sper-
matophores produced by males of this species are huge, representing about 20% of male
body weight (derived from Table 2 in Bowen et al. 1984). Again, females consume the
spermatophylax fully before removing and eating the sperm ampulla. However, the
spermatophylax in this species is almost twice as large as necessary to protect the
ejaculate (Gwynne et al. 1984). Protein from the spermatophylax is incorporated into
Insect Behaviorial Ecology-'85 Quinn & Sakaluk 89
the batch of eggs produced after the mating (Bowen et al. 1984). This suggests that one
function of the large spermatophylax is the production of surviving offspring because it
is likely that the male providing the spermatophylax fathers those young. Gwynne
(1984b) allowed female R. verticalis raised on a low protein diet to eat 0, 1, 3 or 7
spermatophylaxes, then recorded numbers and weights of eggs produced. Volumes of
sperm and other contents of the ejaculate were held constant between treatments. He
found significant increases in both egg weight and egg number produced as a function
of increased numbers of spermatophylaxes eaten. While other benefits to males provid-
ing large spermatophylaxes might exist, the production of surviving offspring may be
important to males. Gwynne (1982) found that when given the choice between two
singing male katydids (Conocephalus nigropleurum), females always mated with the
larger individual. This may represent an adaptive choice since spermatophore size cor-
related positively with male body weight. The potential role that female choice or the
inducement of a refractory period could have played in the evolution of large sper-
matophylaxes remains to be investigated. Indeed, multiple benefits to male reproduc-
tion, some promoting fertilization of eggs and others favoring production of surviving
offspring, appear to be responsible for the large spermatophylax produced by R. ver-
The adaptive significance of benefits provided by males during the prezygotic period
may include increases in both fertilization of eggs and production of surviving off-
spring. The male's cost of providing such benefits is the loss of reproductive opportunity.
The arguments by Alexander and Borgia (1979) identifying such investments as mating
effort ignore the contributions to offspring production.
PREZYGOTIC BENEFITS AND THEIR INFLUENCE ON SEXUAL SELECTION
Under some conditions the benefits to offspring production and survival provided
by males are probably incidental to their attempts to fertilize eggs. For example, mate-
guarding from copulation until oviposition by some odonates presumably evolved to
protect the male's genetic interests by reducing the competition among his sperm and
that of other males. Assuming that benefits of increased fertilization by the male guard-
ing exceed the costs of guarding, the benefits obtained by the female (a period free of
harassment by other males during which she can choose a suitable oviposition site;
Waage 1983) are probably incidental or secondary from the male's perspective, although
this remains to be demonstrated. Because carefully chosen oviposition sites may lead
to increased survivorship of young and because male guarding may enable the female
to take significantly greater care in selecting her site, the effect of this effort by males
could include increased survival of his offspring and reduced female costs for producing
surviving young. In some odonates the method of guarding precludes investment by
males in other mating attempts, and so limits a male's ability to invest in other offspring.
A similar case can be made for mate-guarding that incidentally reduces female predation
risk prior to oviposition, as in the stick insect Diapheromera veliei (Sivinski 1980b,
1983). Assuming the simultaneous existence of other reproductive opportunities for
males, investments of this sort alter the degree to which females limit reproduction,
and thus decrease the competition for mates by males.
The distinction in classification of reproductive effort between the prezygotic and
postzygotic periods suggested by Alexander and Borgia (1979) is misleading when repro-
ductive effort is viewed in terms of its effect on the degree to which one sex versus the
other limits reproduction (sensu Trivers 1972). Male contributions affecting the produc-
tion of surviving young during the postzygotic period usually can be seen easily as
parental effort (Low 1978). We suggest that the main reason postzygotic investments
are easier to understand as parental investment is that in most cases the primary
90 Florida Entomologist 69(1) March, 1986
function is unequivocally for the production and survival of offspring. The primary
function of a prezygotic investment does not alter the investment's effect as limiting
reproduction. Various types of prezygotic and postzygotic investments involving bene-
fits provided by males reveal that irrespective of their primary functions, the impact
on sexual selection's intensity is the same (see Gwynne 1984a). A similar conclusion was
reached independently by Zeh and Smith (1985). Increases in male reproductive effort
that enhance offspring survival may reduce female costs of producing independent
offspring and increase the degree to which female reproduction is limited by the male's
ability to provide that effort. This in turn should lead to reduced intensity of sexual
selection on males.
Male contributions to offspring do, indeed, influence the operation of sexual selection
and affect the courtship roles adopted by male and female insects. In the giant water-
bug. A. herberti, females are aggressive in courtship, although male display is an essen-
tial element of the courtship (Smith 1979b). Sex roles in this species are not completely
reversed, probably because the the costs of egg production and paternal care are equi-
valent (Smith 1979b). When parental investments by males and females are equal,
theory predicts that the sexes should be equally eager to mate (Trivers 1972). Gwynne's
(1981) study of Mormon crickets, Anabrus simplex, showed that prezygotic male invest-
ments can have an equal or greater impact on sex roles in courtship. During mating, a
male A. simplex transfers a very large spermatophore that accounts for up to 27% of
a male's body weight to the female. Females consume the proteinaceous spermatophylax
and then the ampulla after it has been emptied of sperm. Gwynne demonstrated that
females compete for access to males whereas males exhibit mate preference, actively
rejecting smaller, less fecund, females as potential mates.
In some species, males provide "gifts" or services that aid in the production of
surviving offspring. These aids to reproduction can be provided before or after zygote
formation. During the prezygotic period the function of such investments may be exclu-
sively to increase male success at fertilizing eggs. This is more likely to be the case for
discrete adaptations such as a bipartite spermatophore. Continuous traits such as the
size of a spermatophore represent a compromise between the benefits (increased
number of eggs fertilized and increased number of sired offspring surviving) and the
costs (lost reproductive opportunities).
The primary function, or "intention", of male reproductive effort has no bearing on
the operation of sexual selection (i.e., the direction and intensity of the selection).
Instead, what matters is the effect of that effort on the degree to which males versus
females limit reproduction. We suggest that the classification of components of repro-
ductive effort according to the "intentions" of the investor, or the primary function of
the investment, should be avoided. The identification of function is operationally dif-
ficult, and the exclusion of "incidental" effects may result in erroneous conclusions.
Furthermore, we suggest that the original concept of parental investment (sensu Triv-
ers 1972) is most suitable for understanding the relationship between parental invest-
ment and sexual selection.
We thank James E. Lloyd for inviting us to participate in the Insect Behavioral
Ecology Symposium. The Florida Entomological Society and the University of Ok-
lahoma Zoology Department and Graduate College provided travel funds so that one of
us (JSQ) could present the oral version of this paper. We are grateful to Todd A. Crowl,
Insect Behaviorial Ecology-'85 Quinn & Sakaluk 91
Katherine D. Graham, Douglas W. Mock, Patricia L. Schwagmeyer, and John Sivinski
for helpful comments on the manuscript. Randy Thornhill and David Zeh and Robert
L. Smith graciously provided copies of manuscripts that were in press. J. S. Q. was
supported by a Welder Wildlife Foundation Fellowship during the writing of this paper.
Page charges were met, in part, by a NSERC operating grant (4946) to Glenn K. Morris.
1. The concept of parental investment has become obfuscated. Low (1978) divided
reproductive effort into two components, mating effort and parental effort. She defined
mating effort as "any expenditure of nutrient or effort or taking of risks to secure
matings." Parental effort was defined as "any expenditure of nutrient or effort or taking
of risks in the production and raising of offspring or other kin." This divides reproductive
effort according to the function of the effort (for mating or for production and raising
of offspring) and removes the emphasis from the effect of the investment on production
and survival of offspring. Furthermore, effort with multiple functions would be class-
ified according the primary function alone. In this scheme, incidental benefits to offspr-
ing production and survival are neglected. Despite the apparent inconsistency of this
classification with that of Trivers (1972), Low considered parental investment to be that
portion of parental effort received by an individual offspring.
2. Gwynne (1984a) also suggested that postzygotic investment by males could function
as mating effort. Regarding the egg-brooding of male waterbugs, he stated "As long as
there is space available for eggs on a male's back (there is some evidence that male
backs are a limiting resource for females; Smith 1979a), the investment can be consi-
dered ME involved in advertising the male's parental abilities that may aid in the
acquisition of matings with other egg-laden females." There are two problems with this
interpretation. First, if the possession of eggs functions as a signal attracting other
mates, "unattractive" males lacking eggs might be expected to accept eggs without
requiring copulations. Males apparently do not allow oviposition on their backs unless
they have just copulated with the ovipositing female (Smith 1979a, b). Furthermore,
a partially egg-laden male must have been able to obtain at least one mating without
having any eggs on his back. If egg-brooding functioned as a signal of a male's quality,
it is not clear why a male would employ the signal if he were able to obtain matings
without it, especially given its obvious costs. Second, that females exploit information
made available to them by egg-brooding males, thereby allowing females to make opti-
mal mate choices, could be an incidental effect of the behavior (see Otte's 1974 distinction
between the evolved function and incidental effects of 'signals'). Gwynne (1984a) did
concede that paternal care exhibited by A. herberti after the male is completely encum-
bered functions to increase the survival of the young.
3. Because females can store sperm and mate with more than one male, parentage is
less certain for male than for female insects of many species. If benefit-providing males
enjoy increased certainty of paternity over males that fail to provide benefits, then the
degree of paternal certainty will affect the level of investment favored by natural selec-
tion. Whether or not males provide benefits is determined by natural selection according
to the net benefits of available options (e.g., whether to provide benefits, and if so, to
what extent). Males who provide benefits may experience enhanced offspring survival,
and/or increased mate attraction and/or increased probability of fertilization of eggs,
hence increased certainty of paternity (Zeh and Smith 1985). The cost of providing
benefits may be the loss of other mating opportunities. Males that do not provide
benefits may experience increased mating opportunities, but may have reduced offspr-
ing survival, and/or probability of mate attraction, and/or certainty of paternity.
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