Insect Studies in the
Advancement of Science:
P 6 SX8G1 L4
From Darwin and Wallace to the Present
Peter W. Price
Abstract: Studies on insects have provided a wellspring for major advances in ecology and related fields
over the past two centuries. From animal–plant interactions to zoogeography, the insects have inspired
and motivated innovation in science.
Key words: Ecology, historical entomology, evolution
E
cology may be the most central conceptual
field for entomologists. In agriculture,
forestry, and horticulture, plant–insect interactions receive great attention. In forensic entomology, ecological succession is fundamental
because the timing of death is calibrated by the
progress of colonization of necrophorous insects
and the extent of population development on a
corpse. Epidemiology of vector-borne diseases and
the population dynamics of vectors are essential
elements of medical biology. Much of insect physiology is ecologically oriented, and toxicology has
embraced an ecosystem view of trophic relationships along which toxins and their degradation
products may pass. Biological control of weedy
plants and insects emphasize the interaction of
plants and insects in their ecological surroundings.
Regulation of pest numbers encompasses an understanding of predator–prey and parasitoid–host
relationships and population dynamics at several
trophic levels. The evolution and behavior of insects use settings in ecology as an arena for interactions within and among species. Pest management
encompasses many of these subjects in an integrated
manner to maximize the beneficial environments
for crop health and to minimize pest impact. Thus
ecology is central to much of entomological science.
It is for this reason, perhaps, that entomologists have contributed so importantly to the development of basic ecological principles. When we
examine the fundamental contributions that insect
studies have made to ecology, the list becomes impressive and even surprising in my opinion. I emphasize contributions from insect studies because
there is a continuum between those who would call
themselves “entomologists” and those who have
164
not aspired to such a high calling in life—evolutionary biologists, ecologists, and plant protectionists, for example. However, entomologists have a
right to be proud of a rich legacy from insect studies that has generated new areas of ecology and
transformed others.
My aim in this article is to suggest that insect
studies have played a central role in the development of major themes in ecology and related fields—
themes that are largely inseparable from ecology. I
think that most readers will be surprised at the
influence of such studies, as I was when my attention was focused on the issue last year. The occasion was a symposium at the annual national meeting of the Entomological Society of America in Fort
Lauderdale, November 2002. The program symposium, conceived and convened by Carol
Sheppard, was titled “Classic Contributions of
Entomological Studies to Major Biological Subdisciplines”.
Two of the subdisciplines of biology were ecology and evolution. Of course, these areas are inextricably entwined, especially when we enquire as to
why interactions proceed in a certain way, or why
relationships develop in a particular fashion; and
we can readily identify in most major players in the
development of ecology, the ecologist and the evolutionist. Importantly, many of the players were
also bona fide entomologists.
The Naturalist Explorers
Many of the initial players developed during
the age of the traveling naturalists in the 19th century; and whether interested primarily in plants or
animals, most naturalists noted the insects, either
as beguiling or as a nuisance. The epic voyages
were recounted in many books avidly sought by
AMERICAN ENTOMOLOGIST • Fall 2003
the developing middle classes of the day because
nature study and appreciation were a popular avocation (Fig. 1). Alexander von Humboldt and Aimé
Bonpland traveled from 1799 to 1804, exploring
the Orinoco and Amazon Rivers, savannas and
rain forests, collecting 60,000 specimens, and publishing their 30 volumes of Personal narrative of
travels...(1805–1834).
Charles Darwin was greatly stimulated by these
reports and, of course, engaged in his own circumnavigation of the globe from 1831 to 1836. His
escapades and discoveries were reported in The
Journal of Researches (1839), more widely known
today as The Voyage of the Beagle. That Darwin
was an entomologist there can be no doubt. His
enthusiastic collecting of beetles is recounted in his
autobiography (Darwin 1892), including the famous story of the discomfort he experienced after
popping a beetle into his mouth, ostensibly for
safe keeping. Darwin was also an original and life
member of the Royal Entomological Society of
London, becoming vice-president in 1838 (Neave
1933).
Henry Bates acted as president of the same society for 1868 to 1869 and again in 1878, and he
interested Alfred Russell Wallace (Fig. 2) in insects
when they met in 1844. They set out together for
Brazil in 1848, intending to solve the origin of species problem (McKinny 1972). They financed the
expedition with a plan to sell duplicate insect specimens upon their return to London. Bates wrote of
his travels in The Naturalist on the River Amazons
(1863) and Wallace published A Narrative of Travels on the Amazon and Rio Negro...(1853). Wallace
became president of the Royal Entomological Society of London in 1870–1871; his writing desk now
sits in the society’s rooms and is used by chairs of
all meetings (Neave 1933).
Other notable voyages were recorded by naturalists, for example, Reise in Brasilien by J. B. Spix
and C. F. P. von Martius (1824) and Notes of a
Botanist on the Amazon and Andes by R. Spruce
(1908) on travels in the 1850s. Thomas Belt (1874)
recounted his life in Central America in The Naturalist in Nicaragua.
All these voyaging naturalists were amazed by
the insects they witnessed. Darwin thought that
butterflies captured the essence of the tropics more
than any other group and noted flocks of butterflies off shore, such that a mariner cried that “it
was snowing butterflies” (1839, p. 158). Spix and
Martius (1831) observed the strange swollen petioles of melastomes that housed ant colonies; and
Belt (1874) noted ants collecting protein bodies
from leaflet tips of bull’s horn acacias, later to be
called Beltian bodies (e.g., Wheeler 1910).
Emergent Disciplines
Embedded in all this exploration, excitement,
and discovery were the germs of major breakthroughs in ecological and evolutionary understanding. Notably, Darwin (1858, 1859) and
Wallace (1858) formulated the theory of evolution, emphasizing that biotic interactions—that is,
AMERICAN ENTOMOLOGIST • Volume 49 Number 3
Fig. 1. Fascination
with insect life was
popular in the
1800s, reflected in
many illustrations
such as this one, of
cooperation among
insect individuals.
From the
gregarious feeding
of Mourningcloak
butterfly
caterpillars,
Nymphlis antiopa
(L.), to presocial
burying beetles
(Necrophorus spp.)
and the eusocial
wasps and hornets
(Vespidae), all
came under close
inspection by the
wood engraver.
From Dover
Publications (1979)
p. 249.
ecological interactions—were the grist for the evolutionary mill. Darwin repeatedly used insect examples in The Origin of Species because so frequently, they provided particularly revealing illustrations and problems. For example, in the chapter on instinct, he admits to “one special difficulty,
which at first appeared to me insuperable, and actually fatal to the whole theory. I allude to the neuters or sterile females in insect communities” (Dar-
Fig. 2. Alfred Russel Wallace at age 24 in 1848, the
year he sailed with Bates for the Amazon Basin.
From a daguerreotype (McKinney 1972 p. 14).
165
win 1872, p. 218). He went on to discuss “the
acme of the difficulty” concerning caste polymorphism in social insects (p. 220). In addition, the
theory of evolution was the wellspring for new
fields of science initiated or strongly boosted by
the study of insects.
A chronological list of the emergence of new
fields of ecology and their main protagonists will
vary according to personal perceptions. Therefore,
my views are but a series of suggestions that may
foster considerable debate in some cases (Table 1).
Nevertheless, I believe that there is merit in recognizing scientists who have studied insects and
thereby generated new scholarly disciplines in ecology. I will justify some choices briefly.
Although Darwin and Wallace provided a view
of biology from which many derivatives emerged, I
think that the field of population dynamics predated The Origin of Species by many years, but
has remained aloof from evolutionary principles
largely until the present (Price 2003). Records of
forest insect dynamics go back at least to 1800 in
Germany (Klimetzek 1990), and just whom to recognize among all in that great forest entomology
tradition is a matter of choice. My preference is for
J. T. C. Ratzeburg, who in 1840 published his volume on forest Lepidoptera on which long-term
studies had commenced.
The advent of biological control can be traced
to C. V. Riley’s initiative to send parasitoids of the
plum curculio, Contrachelus nenuphar (Herbst),
to other parts of Missouri (Doutt 1964); L. O.
Howard’s (1897) population study of the
whitemarked tussock moth, Orgyia leucostigma
(J. E. Smith), yielded a view of population oscillations generated by the interaction of insect herbivores and their parasitoids. “With all very injurious lepidopterous larvae...we constantly see a great
Table 1. New fields of science initiated or strongly boosted based on the
study of insects
Subject
1. Population dynamics
2. Evolution
3. Ecology
4. Sexual selection
5. Pollination biology
6. Biological control
7. Plant–animal interactions
8. Biogeography
9. Mimicry
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
a
Predator–prey interactions
Ethology
Demography
Ecological genetics
Industrial melanism
Chemical ecology
Coevolution
Sex ratio theory
Tropical ecology
Sociobiology
Biodiversity
Year of Publication
1840
1858, 1859
1859
1871
1862, 1876, 1877
1870a
1875
1876
1862
1879
1897
1914
1921
1930, 1964
1955
1959, 1969
1964
1964a, b, 1967
1966
1975
1988, 1992
No publication in this year, date cited in Doutt 1964, p. 31.
166
Author
Ratzeburg
Darwin and Wallace
Darwin
Darwin
Darwin
Riley
Darwin
Wallace
Bates
Müller
Howard
Frisch
Pearl and Parker
Ford and Ford, Ford
Kettlewell
Fraenkel
Ehrlich and Raven
Hamilton
Janzen
Wilson
Wilson
fluctuation in numbers, the parasite rapidly increasing immediately after the increase of the host species, overtaking it numerically, and reducing it to
the bottom of another ascending period of development” (Howard 1897, p. 48).
This passage probably stimulated A. J. Lotka
(1924) to formulate what came to be known as
his predator–prey equations, the cornerstone of
much modeling of interactions in ecology. Lotka
(1924, p. 90) quoted this passage by Howard
when developing his equations for “a cyclic or
periodic process” of parasite and host population interactions.
Darwin himself initiated several important fields
in ecology (Table 1). Even ecology as a discipline
was fostered strongly in The Origin of Species, for
Darwin repeatedly emphasized biotic interactions
as the most important components of natural selection. Darwin also developed the concept of sexual
selection in The Origin...(chapter 4 on Natural Selection) and in detail in The Descent of Man, and
Selection in Relation to Sex (1871). Pollination biology received much attention from Darwin, and
three of his books (1862, 1876, 1877) established
the field. With these books and his book on insectivorous plants (1875), Darwin also provided the
basis for the large field of plant and animal interactions.
As well as contributing to the theory of evolution, Wallace’s collections of insects and other
groups provided the foundation for biogeography (Wallace 1876). The field was developed further by another entomologist, Philip Darlington
(1957, 1965) whose specialty was carabid beetles.
Henry Bates spent seven and a half years in the
Amazon Basin collecting almost 15,000 species of
insects, of which about 8,000 were new to science.
Based on his field observations and extensive collecting, he developed the concept of mimicry (later
recognized as Batesian mimicry; Bates 1862, 1863),
but only after reading The Origin of the Species
when he returned to England (Gilbert 1983). Fritz
Müller (1879) described what subsequently became
known as Müllerian mimicry (Fig. 3). In fact, Bates
(1862, 1910 edition, p. 348) predicted accurately
the value of studying butterflies for the advancement of science: “The study of butterflies—creatures selected as the types of airiness and frivolity—instead of being despised, will some day be
valued as one of the most important branches of
Biological science” (Fig. 4).
In the early 20th century, other disciplines were
initiated based on the study of the insects. Ethology’s
pioneering spirit must be recognized as Karl von
Frisch, who started his publications on the perception of color in bees in 1914, and Niko Tinbergen
who, early in his career, studied the orientation of
the bee-hunting wasp, Philanthus triangulum Fabricius (Tinbergen 1935, Tinbergen and Kruyt
1938). Karl von Frisch, Tinbergen, and Konrad
Lorenz received the Nobel Prize in Physiology or
Medicine in 1973 for their creative forces in the
field of ethology.
With their studies on survivorship curves for
AMERICAN ENTOMOLOGIST • Fall 2003
Drosophila spp., R. Pearl and S. L. Parker (1921)
brought the field of demography into basic biology, and E. B. Ford (Ford and Ford 1930, Ford
1964) established the field of ecological genetics.
H. B. D. Kettlewell published his first paper on
industrial melanism in 1955. (This topic may be
regarded more as a subdiscipline in a larger field,
such as animal camouflage or animal coloration,
that would necessarily include mimicry. Hence, I
prefer to partition mimicry and melanism to emphasize the important scientists in these areas.)
Two classic papers by G. Fraenkel (1959, 1969)
on the adaptive significance of secondary
phytochemicals played an important role in boosting the study of chemical ecology, although others,
such as Thomas Eisner (e.g., 1970) may well be
rivals for recognition (cf. Sondheimer and Simeone
1970) (Fig. 5). This field was expanded into the
subject of coevolution by Ehrlich and Raven (1964)
using butterfly and plant interactions and further
developed by Thompson (1982, 1994), whose research subjects included swallowtail butterflies and
prodoxid moths. In the same vein of plant and
insect interactions, Janzen’s (e.g., 1966) early papers on ants and acacias in the dry tropics of Central America provided a tremendous boost to the
field of tropical ecology, coupled with his exemplary initiatives in training students in the field. In
the 1960s, Hamilton (1964a, b, 1967) also explained one of the bases of social behavior and
provided a strong stimulus to the advancement of
sex ratio theory.
Understanding of the structure of animal societies was again advanced based on insect studies
by E. O. Wilson (1975) whose expertise in insect
societies (1971) enabled him to create the new field
of sociobiology. Building on the biogeography of
Wallace and Darlington and his own incomparable
knowledge of ants, Wilson (1988,1992) strongly
promoted the field of biodiversity. Wilson also contributed seminally in several subdisciplines, such
as island biogeography (MacArthur and Wilson
1967), character displacement (Brown and Wilson
1956), and the taxon cycle (Wilson 1961).
Fascination and Inspiration
This, then, is an impressive record of how insect
studies have contributed to the development of ecology and related fields. Such a record gives us cause
to wonder why the insects have played such a pivotal role in the advances of biology. But the answers are clear enough to those who study and
admire these remarkable organisms (see box, “Insects as the Inspiration and the Stimulus for Creative Energy and Innovation in Science”). As
Vladimir Nabokov wrote in his autobiography,
Speak, Memory, “Few things indeed have I known
in the way of emotion or appetite, ambition or
achievement, that could surpass in richness and
strength the excitement of entomological exploration” (1966, p. 126).
Insects are, at the same time, endlessly fascinating and puzzling, and solving the puzzles provides
great happiness in one’s existence, according to
AMERICAN ENTOMOLOGIST • Volume 49 Number 3
Fig. 3. Eltringham’s (1916) illustrations of what he regarded
as models and mimics, with races of Heliconius erato and
close relatives as “models” and races of H. melpomene and
close relatives as “mimics.” In fact, both H. erato and H.
melpomene groups are now regarded as members of a
Müllerian mimicry complex (e.g., Gilbert 1983). Heliconius
spp. are now known to sequester cyanogenic compounds
synthesized by Passiflora host plants (Spencer 1988) or
may, in some species, synthesize their own cyanogenic
defensive compounds (Brown et al. 1991).
Linus Pauling. Surely insects must pose more
puzzles than any other group of organisms, being
by far the most diverse and widespread on the earth.
Part of the inspiration and stimulus for study
that insects provide is their visibility and beauty.
No wonder butterflies have indeed been so important in the advancement of ecology, as Bates predicted. The fields in ecology named in Table 1, deriving importantly from the study of butterflies,
include biogeography because Wallace used the distribution of swallowtails to demark biogeographical realms, mimicry (of course), ecological genetics,
and coevolution. Including the moths, we can add
industrial melanism.
Kinds of Scientists
We may then enquire as to the kind of scientist
that is likely to have made the breakthroughs and
conceptual leaps noted in Table 1. My view is that
in each case, scientists had a particularly intimate
167
Fig. 4. Examples of the extreme polymorphism observed in female
Papilio dardanus in Africa, showing various toxic and aposematic
models down the center of the figure and a mimetic morph of P.
dardanus on either side (from de Beer 1964). Where no toxic and
aposematic models exist (for example, on the island of Madagascar),
females resemble the male coloration, as at the top of the figure (male
left, female right).
and extensive knowledge of at least one insect
taxon. A Feeling for the Organism—as Evelyn Fox
Keller (1983) titled her book about Barbara
McClintock—is evident when deep insights are revealed. McClintock discovered “jumping genes”
or transposable elements in corn plants and became a Nobel laureate in 1983.
As Keller noted, “Good science cannot proceed
without a deep emotional investment on the part
of the scientist” (1983, p. 198). But discovering
big patterns in nature, such as those that open
doors to new disciplines in science, requires broad
knowledge of a major taxon or taxa. The scale of
knowledge escalates. Broad comparative approaches are needed in population and community studies and at larger scales. Whereas Barbara
McClintock pondered the thousands of individual
corn plants in a field, a breakthrough in ecology
requires contemplation on thousands of species in
a taxon. Hence, many novel fields have been generated by systematists-cum-ecologists. Just to name
a few:
168
• Edward Wilson, ants and sociobiology and
biodiversity;
• Paul Ehrlich, butterflies and coevolution;
• E. B. “Henry” Ford, butterflies and ecological genetics;
• Sir Richard Southwood, Hemiptera and
Homoptera and insect community richness;
• Guy Bush, tephritid flies and sympatric
speciation;
• Charles Michener, bees and social behavior;
• Howard Evans, wasps, ethology and evolution.
Other great entomologists, including some I
have mentioned already, who showed a deep fascination with their subject include Karl von Frisch
on bees, Gottfried Fraenkel on calypterate flies,
Vincent Dethier, also on flies, and George Varley
on oak-dwelling insects. If we go back to the
founders of our discipline, the list expands almost
endlessly.
Knowing one group of insects intimately provides insights that can be applied more broadly.
The wider the comparative framework, the greater
is the scientific contribution. For example, early
in Edward Wilson’s boyhood, he entered “a magic
kingdom” of natural history (Wilson 1994, p.
47) which, of course, included ants. From a broad
knowledge of ants, he entered into the comparative biology of all social insects (Wilson 1971)
and then to all social organisms in his monumental Sociobiology: The New Synthesis (Wilson
1975). In his autobiography, Wilson noted “But
this is the way it is supposed to be. Nature first,
then theory...Love the organisms for themselves
first, then strain for general explanations, and,
with good fortune, discoveries will follow” (1994,
p. 191).
Educational Involvement
The involvement of those who have studied insects in the development of ecology is further revealed by their contributions to education. One
indicator is the number of general ecology textbooks, including related subject areas, written by
those fascinated by their interactions with insects
(Table 2). The list is impressive. It reinforces the
perception that insect studies and ecology are intimately intertwined. Not all authors listed are considered entomologists, or as being heavily involved
with the study of insects, so some explanations are
in order. I also note the kinds of studies undertaken by other authors better known for their work
on insects.
Although several of the following authors
would not be thought of as working on insects,
their early work was indeed on this taxon. For
example, Elton (1925) published on “The disperal
of insects to Spitzbergen” and soon after on wood
ants, orthopterans, staphylinids, stoneflies, and
mosquitoes (references in Elton 1966). His continued interest in insects is well reflected in his book
on animal invasions (Elton 1958). Shelford’s (1907)
earliest work was on tiger beetles in relation to
plant succession, which may have been a prelude
AMERICAN ENTOMOLOGIST • Fall 2003
to his long-standing interest in ecological succession and communities in North America (references
in Shelford 1963). Some of Hutchinson’s earliest
work was on aquatic insects, including a revision
of corixids in India (Hutchinson 1940, 1959).
Other authors hardly need an introduction as
entomologists or as students of the insects.
Bodenheimer published extensively on the population dynamics of insects, as well as on insects as
food for humans (1951). Among the authors of
the classic Principles of Animal Ecology, Allee studied insects and other arthropods, Emerson the termites, both of the Parks studied Tribolium, and
only Schmidt worked on more depauperate taxa.
In the classic text of the 1950s, Andrewartha and
Birch were both devoted entomologists. So was
Itô.
Schwerdtfeger was one of the great forest entomologists of his time. Williams studied the migration of butterflies initially and moved into
quantitative ecology while still depending heavily
on large insect community samples. Southwood’s
work on insect herbivore population dynamics
and communities is well known. Watt was heavily
involved with modeling insect populations.
Lewontin created his reputation by studying
Drosophila population genetics. Dempster and
Remmert worked principally on insects. Kitching
spent much research time on insects in tree holes
and coauthored a book on insect ecology
(Matthews and Kitching 1984). Putman worked
on blowflies before moving on to vertebrates, and
Wratten has published consistently on insect pests
in agriculture and their natural enemies. As senior author of the well-used text, Begon conducted
research on Drosophila and parasitoids, and Stiling
has published on insects in Spartina salt marshes
and on gall-inducing insects.
I am sure that many other scientists should
be noted as contributing to the development of
ecology based on their experiences with insects,
and I apologize for any omissions, especially to
those who have written in other languages. But
even from my own limited perspective, those who
study insects have made tremendous contributions to the field. Note that Australia, Germany,
Japan, the United Kingdom, and the United States
are well represented in Table 2; and there is every
reason to think that insects will be the centerpiece for further developments. They are, after
all, ubiquitous in terrestrial and fresh water systems and function in important roles as herbivores, animal parasites, decomposers, pollinators, seed dispersers, and, above all, fascinating
subjects to study.
Why insect studies have been so central to the
advancement of ecological science is partially explained in the box (on the next page) and the process of advancement involves initial fascination and
careful study in a comparative way: patterns are
detected, generality is found, and explanations can
be developed. The process was illustrated in the
case of Edward Wilson, the foremost entomologist
of our time.
AMERICAN ENTOMOLOGIST • Volume 49 Number 3
Another example can be seen in the work of G.
Evelyn Hutchinson, who became one of the most
influential ecologists of his time. He studied corixid
bugs in India and revised the fauna (Hutchinson
1940). He noted differences in species richness and
morphology in coexisting species and sought explanations. One result was his hypothesis on limiting similarity in coexisting species, written into his
classic paper “Homage to Santa Rosalia, or, Why
Are There so Many Kinds of Animals?”
(Hutchinson 1959). Perhaps a mammalogist would
be less likely even to conceive the question, let alone
answer it. But one who had studied the insects had
a mass of information with which to tackle the
challenge.
Our challenge is to continue the tradition of the
entomologists and other students of insects who
have forged a pattern of creativity by founding
Fig. 5. Descriptive aspects of plant and insect interactions and predation
on herbivorous insects were well developed in the 1800s, as illustrated
in this wood engraving of nocturnal herbivores and predators. An avid
collector, lamp in hand, appears in the background. From Dover
Publications (1979) p. 253.
169
Table 2. Textbooks in general ecology and related subjects by students of the insects
Year
1. 1927
2. 1938
3. 1949
4. 1954
5. 1958
6. 1959
7. 1963
8. 1963
9. 1964
10. 1966
11. 1968
12. 1968
13. 1968
14. 1971
15. 1973
16. 1975, 1976
17. 1975
18. 1975
19. 1977
20. 1978
21. 1978
22. 1980
23. 1980
24. 1983
25. 1984
26. 1984
27. 1986
28. 1987
29. 1986
30. 1992
Author
Elton
Bodenheimer
Allee, Emerson, Park, Park
and Schmidt
Andrewartha and Birch
Bodenheimer
Itô
Shelford
Schwerdtfeger
Williams
Southwood
Watt
Schwerdtfeger
Lewontin
Wilson and Bossert
Watt
Itô
Schwerdtfeger
Dempster
Ehrlich, Ehrlich, and Holdren
Hutchinson
Itô
Itô
Remmert
Kitching
Putman and Wratten
Andrewartha and Birch
Begon, Harper, and Townsend
Ehrlich and Roughgarden.
Ehrlich
Stiling
Insects as the inspiration and the stimulus for creative energy and
innovation in science
Great diversity of insects provides the strongest comparative power
available
Excellent size range for direct observation in the field and laboratory
Excellent size range for experimental studies
Many species active in daylight
Insects attract attention. Many species are beautiful, dynamic
fliers, with spectacular and diverse design and fascinating behavior
Insect collection is simple and satisfies basic collecting and ordering desires
Serious pests in agriculture, forestry, horticulture, animal husbandry, and
human health
Many economically beneficial species
Ubiquitous distribution in terrestrial and fresh water habitats
All these attributes capture the attention of scientists who often devote a
lifetime to a group. The developed depth of knowledge results in broad
pattern detection and mechanistic explanation of pattern—the basis
for scientific theory
170
Title
Animal ecology
Problems of animal ecology
Principles of animal ecology
The distribution and abundance of animals
Animal ecology today
Comparative ecology (in Japanese)
The ecology of North America
kologie der Tiere, Vol. 1.Autökologie (in German)
Patterns in the balance of nature
Ecological methods
Ecology and resource management
kologie der Tiere, Vol. 2, Demökologie (in German)
Population biology and evolution
A primer in population biology
Principles of environmental science
Animal ecology. 2 vols. (in Japanese)
kologie der Tiere, Vol. 3, Synökologie (in German)
Animal population ecology
Ecoscience: population, Resources, environment
An introduction to population ecology
Comparative ecology (in Japanese)
Comparative ecology (in English)
Ecology: a textbook
Systems ecology
Principles of ecology
The ecological web
Ecology: individuals, populations, and communities
The science of ecology
The machinery of nature
Introductory ecology
new disciplines in science or by invigorating a field
to a new level of recognition. There is no reason to
believe that the creative pace should slacken. The
insects are still out there to puzzle and amaze us
and to stimulate our scholarly impulses.
Acknowledgments
Many thanks are due to Carol Sheppard, who
suggested the topic of this paper and invited me to
speak in her symposium. I am also most grateful to
the Entomological Society of America for financial
assistance in attending the Fort Lauderdale meeting. Rick Karban and Richard Root provided interesting and valuable comments on a former draft
of this paper. I appreciate their expertise and effort
devoted to its improvement.
References Cited
Allee, W. C., A. E. Emerson, O. Park, T. Park, and K. P.
Schmidt. 1949. Principles of animal ecology.
Saunders, Philadelphia.
Andrewartha, H. G., and L. C. Birch. 1954. The distribution and abundance of animals. University of
Chicago Press, Chicago.
Andrewarth, H. G., and L. C. Birch. 1984. The ecological web. University of Chicago Press, Chicago.
Bates, H. W. 1862. Contributions to an insect fauna of
the Amazon Valley. Trans. Linn. Soc. London 23:
495–566.
Bates, H. W. 1863. The naturalist on the River Amazons. Murray, London.
AMERICAN ENTOMOLOGIST • Fall 2003
Photograph is of bee
niches or bee boles:
brick or stone structures in which straw
beehives were placed to
help protect them from
harsh weather. These
particular niches are
over five centuries old
and are found in
Devon, England. The
study of such structures is part of the
archaeology of beekeeping.
WHAT IS IT?
ANSWER:
AMERICAN ENTOMOLOGIST • Volume 49 Number 3
free illustrations of mammals, birds, fish, insects,
etc. Dover Publications, New York.
Ehrlich, P. R. 1986. The machinery of nature. Simon
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Peter W. Price is Regents’ Professor Emeritus at
search involves the ecology of plant, herbivore, and
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1997).
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