IN MEMORIAM

Ron Prokopy 1935–2004

Ron Prokopy died at his home in Conway, Massachusetts

on 14 May 2004, at the age of 68.

His research career focused on the behavioural ecology

of tephritid fruit flies, particularly the apple maggot fly,

Rhagoletis pomonella, and a spectrum of other insects,

most of them pests on apple or other fruits. His work also

explored the theoretical underpinnings and practical

boundaries of producing apples with a minimal input

of pesticides. This he practiced in Massachusetts in experi-

mental orchards, in cooperation with the community of

Massachusetts apple growers, and in his own orchard in

the Berkshire Hills.

Ronald John Prokopy was born in Danbury, Connecticut

on 28 September 1935. He grew up on his family’s farm in

Danbury, which had various fruit trees including apples.

This experience seemed to provide an imprint for a life-long

passion to devise practical methods for farmers to decrease

reliance on insecticides. He received two degrees from

Cornell, a B.S. in Agriculture in 1957, followed by a PhD

in Entomology in 1964 under the direction of George

Gyrisco. Ron’s dissertation centred on the migratory

behaviour of the alfalfa weevil.

In 1964 he joined The Connecticut Agricultural Experi-

ment Station as an Assistant Scientist in the Department of

Entomology, with a responsibility to improve control

measures for apple pests. The apple maggot was the key

apple pest in the north-eastern United States, dictating a

bi-weekly cover spray of insecticides, from petal fall in late

spring until summer’s end. There his life-long fascination

with the apple maggot fly began. Ron’s initial work

explored the visual ecology of resource location: how the

shape, colour and size of fruit-like objects and foliage

mimics served adults in the location of food and sites for

mating and oviposition. His colleague Jim Kring was

studying parallel questions with aphids and served as an

influential mentor. A hallmark of Ron’s approach to

generating hypotheses and to developing simple but diag-

nostic experimental tests was spending many hours in the

field, simply observing an insect’s movement patterns and

behaviours. Over the next 40 years nearly all of his

behavioural studies would involve day-active insects,

which he could observe in nature.

Ron’s scientific experiences between 1968 and 1975 were

peripatetic. He held Visiting Professor posts at the Pomol-

ogy Institute in Skierniewice, Poland, in 1968 as well as at

the Swiss Federal Research Station in Wadenswil in 1969.

From 1969 to 1973 he served as a Research Associate at the

University of Texas, where he collaborated with Guy Bush.

Guy’s interests in sympatric speciation in Rhagoletis flies via

host race formation, and Ron’s interests in the mechanisms

of host selection dovetailed into studies of this fly’s court-

ship and host-selection behaviours. In 1973–1974, Ron

worked as a UN-FAO consultant in the ‘Democritos’

Research Center in Athens. In 1974 he set up the Prokopy

Bio-Experimental Farm in Bailey’s Harbor, Wisconsin,

where he and many visiting colleagues continued their

field observations of Rhagoletis.

In 1975 Ron joined the Department of Entomology of the

University of Massachusetts, Amherst as Associate Professor,

becoming a Professor in 1981. His responsibilities then

were split between research, teaching and extension. Ron

mentored many graduate and postdoctoral students and

taught a rigorous graduate course on Integrated Pest

Management. His extension responsibility was to devise

pest management programmes for the Massachusetts

apple growers and to encourage their adoption. Ron

Physiological Entomology (2004) 29, 489–490

# 2004 The Royal Entomological Society 489

produced a steady stream of recommendations, which

progressively lowered insecticide use. Every year Ron

would present a dozen ‘Twilight Talks’ to growers around

Massachusetts, persuading this generally conservative com-

munity to adopt his recommendations. That Ron himself

was a grower did much to win over sceptics. His own

orchard served as a crucible for innovation; in recent years

the only insecticide sprayed there was for plum curculio, and

the apple maggot was controlled entirely by being inter-

cepted on a peripheral barrier of visual and odour mimic

of apples laced with a little insecticide. Although his

orchard was isolated from other sources of most apple

pests, an old untreated apple tree in front of the Prokopy’s

farmhouse served as the source of inoculums for apple

maggots and other pests.

Ron’s nearly 30 years at the University of Massachusetts

provided a stable base for his studies of apple and fruit-

infesting insects. Many scientists visited his laboratory for

extended stays and his colleagues in Amherst shared his

interests in behavioural ecology. Vince Dethier especially

served as a source of information on the repertory of fly

behaviours. One of Ron’s cherished activities was to

consider precisely why insects behave as they do. You

would have the clear impression that Ron, while closing

his eyes, would try to envisage himself as the insect perceiv-

ing the very situation you just described to him.

Over the course of his career Ron published 275 referred

journal articles, over 30 major reviews (including four in

the Annual Review of Entomology), and hundreds more

proceedings, extension articles and notes. Among some of

his discoveries with Rhagoletis was the host-marking

pheromone, which is laid down by the ovipositor dragging

on the fruit following oviposition, and which informs later

arriving females that an egg has already been deposited in

the fruit. Subsequently, he demonstrated the role of experi-

ence (prior contact) in females reacting to the mark; that

males tended to increase residence time on a marked fruit,

thereby improving their chances of encountering a female

landing on an apparently favourable fruit; and that the

mark aided host finding by a wasp parasitoid of the fly.

These intricacies exemplify how Ron’s keen sense of obser-

vation could lead to previously unimagined behaviours.

Among the many honours Ron received were a Guggenheim

Fellowship, a Fulbright, and three major awards from

the Entomological Society of America: the Buzzart Award

for Outstanding Contributions to Agriculture, the Foun-

der’s Memorial Award and Lecture (which honoured John

S. Kennedy, another scientist who influenced Ron’s work),

and the Distinguished Award in Extension Entomology.

Ron was characteristically modest about these and many

other awards.

Visiting Ron and his wife Linda at their farm high in the

Berkshire Hills was always a memorable experience. An

afternoon walk through the orchards, interspersed with

Ron’s visions for growing apples without any pesticides,

would be followed by a dinner animated with lively discus-

sions of science, politics, social justice, and stories of Ron

and Linda’s very entertaining life adventures. Many friends

and colleagues will remember forever those evenings at the

Prokopy’s dinner table. All who knew Ron marvelled at

his energy, and his sudden passing therefore was most

unexpected. Ron is survived by Linda, his sons Josh and

Max, and a granddaughter, Annabel.

RING T. CARDE

University of California, Riverside

California, U.S.A.

JOHN G. STOFFOLANO JR

University ofMassachusetts, Amherst

Massachusetts, U.S.A.

490 In Memoriam

# 2004 The Royal Entomological Society, Physiological Entomology, 29, 489–490