Quo Vadis, Taxonomy?l

Quo Vadis, Taxonomy?l
By Curtis W. Sabrosky2
Systematic Entomology
Laboratory, U.S. Department of Agriculture, Washington, D.C.
some data are available. In North America, the largest
concentration of taxonomic entomologists is in Washington,
D.C.: 42 active professionals (31 U.S. Department of
Agriculture
and 11 Smithsonian Institution),
besides
administrators and a few research associates and collaborators. This group is second in the world only to the 59
taxonomic entomologists centered at the British Museum
(Natural History) in London (41 Museum, 12 Commonwealth Institute of Entomology, and 6 of other agencies).
Other large staffs in North America include 26 at the
Entomology Research Institute in Ottawa, Canada, 10 in the
California Department of Agriculture at Sacramento, 6 in the
illinois Natural History Survey at Urbana, and 5 in the
Florida Department of Agriculture at Gainesville.
Tabular comparison is only partially successful, because
records are kept in different ways, by lots received, by
identifications, or by specimens. Table 1 summarizes the
work loads of professional insect taxonomists in the organizations just mentioned.
Without depreciating in the slightest the contributions of
other organizations, I must point out the enormous burden
carried by the insect taxonomists of the U.S. Department of
Agriculture. For many years they have received for identification more than 30,000 individual lots each calendar year. The
number of identifications, or separate names furnished to
correspondents, has averaged nearly 82,000/year for the past
10 years. Specimens identified, exclusive of scale insects,
total more than 322,000 annually. I marvel that in spite of
this burden, the USDA taxonomists have produced many
research papers, notable catalogs on Hymenoptera, Diptera,
termites of the world, and mosquitoes of the world, and
books on special topics such as blow flies, leafhoppers, and
acarology.
However commendable their record, it is not without
fault. The burden and constant pressures often lJIean delays
and disappointing time lags in identification and reporting.
Far more serious is the depressing effect on the research that
should be carried on by those competent specialists. It is a
pitiful fact of life that able specialists wear away the best
years of their lives trying to cope with that ceaseless
flood tide that beats upon their doorsteps.
Many, if not most, requests for identifications are justified,
though not necessarily urgent. No matter what his affiliation,
no taxonomist should need to procrastinate or refuse. If tasks
are justified, then the task force should be commensurate in
size and support with the need and the demand. And for
taxonomists in general the justifiable demands are escalating
with the increase in such projects as environmental studies,
biological control activities, IBP, and a sea-level canal.
Quo vadis - where goest thou - taxonomy?
Behind this question lie many others. What is the task of
taxonomy? Is the support commensurate with the task? Do
taxonomists deserve support? Have they, or have they not,
served biology in general, entomology, and their fellows?
Much of my discussion refers to taxonomy and taxonomists in general. Other parts apply particularly to taxonomic
entomology in the Federal Government.
THE TASK OF TAXONOMY
The task of insect taxonomy encompasses the identification ·and classification of
insects
and
related
arthropods, with all relevant activities. For this
audience I need not dwell
upon the immense size and
diversity of the field. There
are hundreds of thousands
of known species, classified
in tens of thousands of
generea and subgenera, organized into thousands of
families and other groups,
with numerous synonyms
in all echelons, all forming
a stupendous agregation of
names and taxa. Yet it
seems safe to say that no
Curtis W. Sabrosky
more than half, possibly
only one quarter, of the
world's insect fauna is known. Perhaps one or two million
species remain to be described, besides which only a small
fraction is known of the immature stages of the already
described species. Obviously a huge task remains to be done.
But taxonomists are not merely inventorying the fauna.
Many new species are only byproducts of more precise
discrimination of the elements of species complexes, sibling
species, cryptic species, and convergences.
At a practical level, the task of taxonomy is reflected by
the service identifications being done by taxonomists individually and in various public agencies in addition to - or
at the expense of! - their research programs. The volume of
service identifications and the associated consultation provided by taxonomists are measures of the task of applied
taxonomy. Such identifications are often vital to the research
of others, and performed as a service by the taxonomist, an
application of his specialized knowledge. For public agencies,
1 Presidential address delivered at the opening session of
the annual meeting of the Entomological Society of America
in Chicago, Dec. 1, 1969.
21 acknowledge with thanks the prompt and generous
cooperation of the following in supplying information on
service identifications and personnel of their organizations:
W. B. Mountain, E. C. Becker, and J. J. H. Martin, Canada
Department of Agriculture; George T. Okumura, California
Department of Agriculture; H. A. Denmark, Florida Department of Agriculture; Philip W. Smith, illinois Natural History
Survey; Maynard J. Ramsay, Plant Quarantine Division, U. S.
Department of Agriculture; Paul Freeman, British Museum
(Nat. Hist.); R. G. Fennah, Commonwealth Institute of
Entomology .
THE WORK OF TAXONOMISTS
Have taxonomists done a good job? In seeking an answer
to such a question, I shall discuss briefly some facets of the
work of taxonomists.
Nomenclature.-Complaints
about changes in familiar
names are frequently heard. A correspondent wrote me this
year that, "It is almost impossible for those of us who are
general entomologists and general biologists to keep up with
all of the changes in names of organisms - to say nothing
about the reasons for those changes."
Name changes may be for zoological as well as for
3
Table I.-Major North American centers for insect identification.
Organization
Systematic Ent. Lab.,
USDA
Dept. Entomology,
Smithsonian
Ent. Research Inst.,
Canada Dept. Agr.
Insect Taxon. Lab.,
Calif. Dept. Agr.
Div. Plant Industry,
Fla. Dept. Agr.
Ill. Nat. Hist. Survey
Taxonomists!
1959-69
Lots
received
No.
identifications
31,3922
81,7432
22
30
5
11
21
26
8352
6,5203
7
10
13,8474
24,6002
5
5
12,0905
5
6
Specimens
identified
322,4282,7
3,5132,8
7,6292,8
42,3852
35,4266,9
8752
~Administrators not included unless also practicing specialists.
3Average of 10 years, 1959-68, inclusive.
4Average of 4 years, 1965-68, inclusive.
5Average of 7 years, 1961-67, inclusive.
6Average of 5 fiscal years, July 1, 1964-June 30, !969.
Based on half-year count of 17,713 for January through June 1969, but because spring is the heaviest period for
id~tifications, the final total for 1969 will probably be somewhat less than estimated total.
8 Not counting scale insects.
From lots received by USDA, or mixed lots received at the Smithsonian Institution and referred to USDA. The figures do
not include lots received at the Smithsonian and exclusively for Smithsonian personnel, for which exact figures are not
available.
9 Counting scale insects.
nomenclatural reasons. Zoological changes result from shifts
in classification or identification, sometimes made more
annoying by differences of opinion among taxonomists.
Depressing as it may sound, many changes are yet to come,
because of the lag in modern revisionary work from the
relatively small number of active taxonomists. Changes for
zoological reasons can usually be satisfactorily explained and
justified. However, changes for purely nomenclatural reasons,
i.e., application of the International Code of Zoological
Nomenclature, somehow never seem so justifiable, no matter
how easily explained. The Law of Priority, which requires the
use of the oldest proposed name that satisfies certain
generally accepted criteria, is a method for complete objectivity in the choice of names, avoiding personal, nationalistic,
or other prejudices. Unfortunately 2 serious limiting factors
exist: (1) the inadequacy of catalogs; and (2) human nature,
the unwillingness to change, to lose face by retracting a
decision once published. The 3 little words, "I was wrong,"
are not often heard in taxonomy. One is more likely to hear,
"I was robbed'"
Rules of nomenclature have no force or enforcement
except the weight of their own merit backed by general
acceptance and world opinion. It is a safe bet that no one
agrees with all rules in the Code, but in general this Code is
what most of us abide by. I am also fIrmly convinced,
contrary to the ideas of individuals who go their lone way,
that stability and universality would be greatly advanced by
united support of a set of rules, rather than by disunity and
the contemporaneous
application of different rules. In
nomenclature I maintain that the end is more important than
the means. Independence of action hinders rather than helps
when we are actually striving toward a common goal - stable
and universal names.
Common Names.- The Society's approved lists of
"Common Names of Insects" include their scientific names,
and in this arena the work of taxonomists is in plain sight of
everyone. Regardless of whether one approves of lists of
common names - and I am aware that some taxonomists do
not - an analysis of the lists and changes in them should bear
witness to the work of taxonomists.
The fust such approved list, published in 1908, consisted
of only 142 pairs of names, common and scientific. This
sample is small, and the scientific names do not appear to
have been critically checked by specialists. The second list
(1925) offers the sizable sample of 541 pairs (or 531,Ieaving
out 10 dropped from later lists), and accordingly this list was
used as my base line for analysis. It includes, in general, the
most common and most important species, on the lists for a
full 40 years, to and including the latest list (1965).
Table 2.-Common-name lists, 1925-65.
(Nine lists.)
Changes
Common names
Scientific names
Totals
25
278
303
Alterations
Totals
III
425
536
136
703
839
Table 2 summarizes the history of the 531 names. There
have been 25 changes and III alterations in common names,
and 278 changes and 425 alterations in scientific names. In
other words, scientific names show 11 times as many changes
and 4 times as many alterations. What do these facts mean?
Should one dwell upon the apparent instability of scientific
names compared with common names? Or should one
deplore even 25 changes in common names as unfortunate
changes of mind? Or should one emphasize above all that 296
(55.7%) of the pairs of names have had no change whatsoever, or only trivial alterations? Clearly we must look
beyond the bare figures.
Alterations are mere technical or trivial details, often based
on editorial policy, that do not affect the essential recognizability of names, such as hyphenation, parentheses around
authors' names, or spelling changes such as the successive I,
11, m, and mm variants of hellgrammite. The great majority
of these came in the first 3 or 4 lists, when there was a
certain amount of indecision on format and apparent lack of
careful checking of details.
4
Table 3.-Changes in scientific names from the 1925 list.
1931
1937
1942
1946
1950
1955
1960
1965
Totals
Generic
name
34
18
35
24
5
15
4
24
159
Specific
name
12
4
23
10
5
18
12
15
99
5
2
8
1
1
2
0
51
24
66
35
11
35
16
Both
Totals
20
40
278
Table 4.-Screw-worm.
What of the really critical point, the numerous changes in
scientific names? Table 3 analyzes these changes, showing
159 in generic name alone, 99 in specific name alone, plus 20
in both at once, and shows that changes are still continuing at
a fairly steady pace. If this condition seems bad, please
remember that scientific names are not mere captions for
pigeonholes. The generic name represents a fllst step in
placing the specific taxon in the hierarchy of zoological
classification, but this step is subject to differences of
opinion among taxonomists. Extensive reclassifications have
resulted from studies of genitalia, host relationships,
immature stages, and cytology. By and large, these are the
causes of the large total of 179 changes in generic names.
Moreover, taxonomy has always been understaffed for the
overwhelming diversity that it faces. Many, many groups
waited years for revision, and many still await modern study.
Incidentally, long overdue reclassification in aphids and scale
insects, especially mealybugs, caused most of the recent
changes.
At the species level, the 119 changes have come about in
various ways, for example, from revisionary studies (as in the
bark beetles, Dendroctonus), from separation of New and
Old World species (Heliothis zea from H. armigera), or from
application of rules of nomenclature (Lygus Iineo/aris, an
older name than L. oblineatus).
Date
Name
1908
1925-31
1937-42
1946-50
1955-60
1965
Chrysomyia macellaria (F.)
Cochliomyia macef/aria (F.)
C. amen'cana Cushing & Patton
Callitroga americana (Cushing & Patton)
C. hominivorax (Coquerel)
Cochliomyia hominivorax (Coquerel)
realized that 2 distinct species were being confused. Aubertin
and Buxton in England adopted hominivorax in 1934, only a
year after americana was named, and they were followed by
the rest of the world, as we, too, should have done.
Table 5.-Margined blister beetie.
Date
Name
1908
1925
1931-42
1946
1950 to date
Epicauta marginata (F.)
E. cinerea marginata (F.)
E. marginata (F.)
E. solani Werner
E. pestifera Werner
Time does not permit a detailed review of name changes.
However, it may be illuminating, after the questions asked
The case of the margined blister beetle (Table 5) is
about the work of taxonomists, to note a few of the
straightforward, bad as it looks. Fabricius' name marginata
sequential changes that I am sure have annoyed many people.
truly applies to a distinct species from the Cape of Good
Other examples might be discussed with profit and interest,
Hope, but our usage dated from an error by the great man
such as the scientific names for the codling moth and the elm
himself, who applied the name marginata to a similar
leaf beetle, but I shall reserve those for later review in a paper
American species. Our species was fmally christened in its
dealing with the common-name lists.
own right, but unfortunately the name chosen, solani, was
The screw-worm (Table 4) illustrates both zoology and
found to have been used previously in another part of the
nomenclature. In 1915, Townsend separated the New World
world, hence the final change, which its author wistfully
forms as Cochliomyia, and in 1933 the confusion of primary
hoped would be the last!
and secondary screw-worms was sorted out. So far, all is
The hornworms (Table 6) show a variety of changes:
legitimate zoology. The rest is a mixture of nomenclatural
Backing and f1lling on the generic name, and most astonishanomaly, human nature, and taxonomic error. Suffice it to
ingly a reversal in the use of the specific names. The generic
say that hominivorax was clearly described and figured over a
name Manduca is older than Protoparce by a half century,
century ago from a case of human myiasis, and should have
but its adoption was delayed by difference of opinion on the
been recognized from the original publication, once it was
acceptability of a certain publication.
Table 6.-Hornworms.
Date
Name
Genus
1908
1925
1931
1937
1942-60
1965
tobacco horn worm
Phlegethontius
Protoparce
Phlegethontius
Protoparce
Protoparce
Manduca
quinquemacula ta
quinquemaculata
quinquemacillata
sexta
sexta
5
tomato horn worm
sexta
sexta
sexta
sexta
quinquemaculata
quinquemaculata
Table 7 .-Spruce budwonn.
Date
1865
1925-31
1937
1942-46
1960 to date
favor of concentration on more important, more needed, and
more useful projects. I suggest that to the familiar expression,
"oh my" insects, we might add another term, "so what"
taxonomy.
Let us be realistic. Today vast sums are involved in poverty
programs, space research, wars past and present, and so on. I
see no immediate hope of any great leaps forward in the
support of taxonomy, federal or otherwise. I may be unduly
pessimistic, but holding that view I favor purposeful selection
of what is studied, cold-blooded determination of what is
important, and concentration on getting those tasks done.
Otherwise I fear that we may end with detailed knowledge of
Micronesia or Malaya or Malawi, but with gaps in our
knowledge of Main Street, U.S.A. We may have revisions of
minor families or rare genera but fail to understand important pests and parasites and predators. I believe strongly
that applied taxonomists need to revise their aims and
priorities and realistically restructure their efforts.
Name
Tortrix? fumiferana Clemens
Harmologa fumiferana (Clemens)
Cacoecia fumiferana (Clemens)
Archips fumiferana (Clemens)
Choristoneura fumiferana (Clemens)
The spruce budwonn (Table 7), my last slide, shows one
consistent specific name but 5 different generic names. The
reasons for change lie almost entirely in successive reclassifications of the tortricid moths, with more restrictive genera
now being recognized.
In general, explanations can be made for many of the
name changes. Some change is inevitable with developing
classifications, and there must be tolerance for legitimate
difference of opinion on such matters as generic position,
specific synonymy, and subspecific status. Yet I must confess
to a nagging feeling that some cracks in the facade cannot be
satisfactorily explained or defended. I cannot escape the
feeling that not all taxonomists have reviewed the lists
carefully and faithfully, and that some errors have been
perpetuated by this laxity, or by fealty to familiar usage. If a
worthy job is to be done, and it is worth doing on a work of
reference and standard, then it is worth doing well.
Having said that, I shall close this section with mention of
a case, not involving a name change, that is defmitely to the
credit of taxonomists, even though they have been charged
otherwise.
Several years ago, at an annual meeting of the Society, a
speaker stated that the seriousness of an immigrant species,
the Japanese beetle, "was masked for years by an entomologist who misidentified the specimens." In reality, the
beetle, even though exotic, was not only identified correctly
and promptly, but a warning of its potential seriousness was
issued by the taxonomist. The beetle was discovered in a New
Jersey nursery in August 1916. Sometime in the first half of
1917 it was identified as Popilia japonica by H. S. Barber in
Washington. Barber warned, according to the collectors,
Dickerson and Weiss, "that this was the first record of the
genus from America, and further that species in allied genera
have caused considerable trouble in the Old World and when
introduced into various of the Pacific Islands."
Dickerson and Weiss promptly revisited the nursery to
make further observations. In other words, within only one
year, the Japanese beetle was discovered, submitted for
identification, identified, reported, and rechecked at the
original site! Less than one year later, the story was in print
in the Canadian Entomologist.
A few general remarks on the work of taxonomists: they
are frequently preoccupied with far-off faunas. A great deal
of work needs to be done on North American insects,
particularly revisory work, catalogs, and handbooks. Yet
some taxonomists and some important institutions emphasize
studies of far-off faunas to the neglect of our own. Studies of
the distant are not all bad, of course. American interests are
broad, high-speed transport brings us exotic species, and
threatened ecological niches invite and demand exploration
before they disappear. Research is research, and defensible as
such, within reason, but if that needed at home is neglected.
then the public and the entomological profession have, in my
opinion, a legitimate reason for questions.
Taxonomists often seem to have a predilection for the
unusual and the intriguing in minor groups of little or no
importance. Such studies have some values, of course, but
when resources of men, money, and time are more or less
rigorously limited, the minor groups should be stockpiled in
THE IMAGE OF TAXONOMISTS
What of the image of taxonomists? Are they old-fashioned,
not with it?
Taxonomists in general are dedicated individuals, sometimes too much so as many a wife can testify. Some of the
dedication admittedly is selfishness and introversion, but
with taxonomy lacking the support that flows to more
popular and dramatic disciplines, dedication has made
possible much of what we know in taxonomy today. This
dedication undoubtedly contributed at one time to the image
of an entomologist as a wild-eyed professor chasing a
butterfly across flowered fields. This image has changed
considerably in modern times, but some of the old image still
remains in cartoon and story. With all the derision, there may
be a touch of envy or wistfulness. The things that taxonomists do reflect that of which they can be proud: enthusiasm,
dedication, and love of one's life work. No apology is needed
for those. Until insect taxonomists draw their final breaths,
they will remain fascinated students of one of the world's
most important groups of animals.
The attitudes of taxonomists are as varied as human
nature. Some have made a career of being good cooperators,
and some a career of being s.o.b.'s. Such a range may exist in
all professions, but perhaps some of the worst prima donnas
are in taxonomy, because taxonomists tend by nature to be
introspective and self-centered. It is probable that the actions
of some of us taxonomists have contributed to our lack of
support. We need somehow to renovate the image of
taxonomists, as well as of taxonomy.
THE SUPPLY OF TAXONOMISTS
Taxonomists are the unfortunate beneficiaries - if I may
use that tenn figuratively - of a modem trend in biology. As
Dobzhansky expressed it a few years ago: "At present,
molecular biology is at the top, and systematics at the
bottom, of the peck order in biology." Originally the
cataloguing of the fauna and flora was the exclusive interest
of biologists. They were impressed by the variety and
grandeur of nature, and they undertook to describe and name
the organisms. Then followed successively the rise of other
fields - comparative anatomy, physiology, genetics, and so
forth - and molecular biology is the latest of these. The tide
moved away from taxonomy, and fmally even against it.
However,
many disciplines, such as entomology,
nematology, and malacology, still have enonnous taxonomic
problems. Unfortunately, taxonomy is in serious need of
competent taxonomists. The deficit is becoming acute at a
time when active fields, such as ecology, need the services of
taxonomists even more. The present taxonomic community
6
is aging, with too few replacements. There is less recognition,
or none at all, in some graduate schools, of taxonomic
problems. I am deeply concerned about the present support
of the training and encouragement of taxonomists for the
future.
Within systematics itself, the status of taxonomy proper is
not high. Some might say sardonically that this condition
stems from the prestige and impact and grant value of key
words like evolution, speciation, and phylogeny. Nevertheless, and whatever the cause of the status, accurate
identification is as important as it ever was, and with the
discovery of complexes of species and large numbers of still
undescribed species, precise discrimination is becoming in
many cases more and more difficult and time consuming.
Furthermore, there is that tendency for taxonomists to
choose glamour insects, the "oh my" groups rather than the
"oh hell" groups. How many study scale insects, aleyrodids,
wireworms, or tachinids, to mention a few highly important
but unpopular groups?
An experienced taxonomist is developed over a period of
time; he does not come Ii la cafte as "instant authority."
Somehow we must replenish and increase our supply of
taxonomists, and concomitantly revitalize taxonomy itself.
to space exploration, lasers, and similar striking developments.
Over the past 2 years, the role of systematics in the
Federal establishment has been reviewed by a Panel on
Systematics and Taxonomy (POST) established by the
Committee on Environmental Quality of the Federal Council
of Science and Technology. The panel's report will be
published in the near future. This year a committee
established by the National Research Council has been
reviewing a proposal for an institute for the support of
applied systematics. In the United States Department of
Agriculture, new insect taxonomists have been added in each
of the past 2 years, so that at present the Insect Identification Unit has 30 specialists, the largest number in its history.
The Plant Quarantine Division has developed a corps of 20
identifiers in various ports and border areas. These identifiers
serve the immediate needs of their organization and materially decrease the pressure on the Washington taxonomists.
Several States, notably California, Florida, and Illinois, have
important centers for taxonomic service work, and in recent
years some other States have sought to develop or increase
their in-house competence in taxonomic work. All this is
helpful and hopeful. We should renew our efforts for total
support commensurate with the task.
Perhaps it is appropriate here to mention the subject of
publications. I look forward to the day when taxonomy,
along with other fields, can fully utilize modern storage and
retrieval systems. It is not yet clear in what direction or how
far we should go with these methods, but we cannot stand
still, or turn our backs on thc problem. As the late President
Eisenhower so well expressed it, "We cannot face the future
simply by walking into the past backwards."
THE SUPPORT FOR TAXONOMY
Is thc support for taxonomy commensurate with the task?
On balance, and with due acknowledgment of taxonomists' failures and weaknesses, I must answer "No." I am
afraid that not all taxonomists do or have done as good a job
as they might, but even if all do a good job, they would still
be swamped. Taxonomists do lack enough material support,
in spite of years of oral support. The hard-working and
conscientious taxonomists are overworked. They deserve
much more help. Personnel of service organizations should be
adequate to do a good service job and to carry on research.
Both are important, and both are essential. In the long run,
doing a good service job demands supportive research, but
research is manifestly difficult under the deadly workload
carried by the USDA taxonomists.
A great
deal more
support
is needed
CONCLUSIONS
We are living, in many respects, in a time of ferment. I am
not so "square" as to propose a rededication to the three r's
- though some of that is badly needed! Rather I suggest
many r's. To review briefly the recommendations in my talk
today, I believe that taxonomists (and, where appropriate, all
entomologists)
at all levels in both
should
spirit and funds. We need much more for the taxonomic
services performed by the overworked Federal taxonomists.
We need increased State support, because there is much that
the States can and should do for themselves locally. We need
more individual cooperation with S~ate and National agencies
in taxonomy. We need more mobilization and coordination
of existing manpower and facilities, because, realistically, the
prospects for any immediate major improvement seem rather
dim, or at least distant.
There are some hopeful signs. The present climate of
opinion is favorable, in spite of the understandable attention
First, revise aims and priorities realistically, and restructure
the effort in applied taxonomy;
Second, renovate the image of taxonomists;
Third, replenish the supply of taxonomists, and concomitantly revitalize taxonomy itself;
Fourth, renew efforts for support commensurate with the
task of taxonomy;
And finally, reaffirm faith in taxonomy.
Revise. Restructure.
Renovate. Replenish. Revitalize.
Renew. And finally, reaffmn.
Quo vadis, taxonomy? Quo vadis?
MATING OF THE HONEY BEE
that worker bees had to be excluded from the cage because
they clustered around the queens. The exclusion of the
workers presented the problem of maintaining drones and
queens without direct contact with workers. The primary
question, if queens would continue mating flights if they had
contact with workers only through a screen, was answered
affllmatively. The queens also mated when these experiments
were carried on outside the cage.
Once the queens and drones were in the cage, the drones
pursued queens as well as other drones. The present problem
is in getting the drones to focus all attention on the queen
rather than on one another.
JOHN HARBO and ROGER MORSE
To selectively breed any animal, man must be able to
control the mating. Honey bee mating is especially difficult
to control because the queen bee flies alone from her hive
and mates with drones which may live miles away. The
methods presently used to control honey bee mating are (1)
stocking all the bee hives for miles around with desirable
drones or (2) artificial insemination. Artificial insemination
gives better control, but it is time consuming and requires
special skill and equipment. If bees could be naturally mated
in cages, controlled mating would be greatly simplified.
Harbo and Morse studied the mating behavior of honey
bees with the hope that it will eventually lead to the ability
to induce mating in confinement. It was quickly established
Department of Entomology
Cornell University
7