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I.  The
recorded history of biological control may be considered as dating from
Egyptian records of 4,000 years ago, 
where domestic cats were depicted as useful in
rodent control.
 
II.  Insect Predation
 
A. 
Recognized at an early date, but the significance of entomophagy and
exploitation was lost except for a 
      few
early human populations in Asia where a sophisticated agriculture had
developed.
 
B.  The Chinese
citrus growers since at least 400 BC placed nests of predaceous ants, Oncophylla 
      smaradina, in trees
where the ants fed on foliage-feeding insects. 
Bamboo bridges were constructed to 
     
assist the ants in their movements from tree to tree.
 
C.  Date
growers in Yemen went to North Africa to collect colonies of predaceous ants
which they 
     
colonized in date groves to control various pests.
 
III.  Insect Parasitoidism
 
A.  Not
recognized until the turn of the 17th Century.
 
B.  First
record attributed to the Italian, Aldrovandi
(1602).  He observed the cocoons of Apanteles 
      glomeratus being attached to larvae of Pieris rapae
(the imported cabbageworm).  He
incorrectly 
     
thought that the cocoons were insect eggs.
 
C.  Printed
illustrations of parasitoids are found in Metamorphosis
by J. Goedart (1662) <PHOTO>. 
      He
described "small flies" emerging from butterfly pupae.
 
D.  Antoni
van Leeuwenhoek in 1700 (van Leeuwenhoek 1702) described the phenomenon of
parasitoidism 
      in
insects.  He drew a female parasitoid
laying eggs in aphid hosts.  
 
E. 
Vallesnieri (1706) <PHOTO>, first correctly interpreted this
host-parasitoid association and probably 
     
became the first to report the existence of parasitoids.  Bodenheimer (1931), however, noted that
several 
     
earlier entomologists recognized the essence of parasitoidism.
 
F. 
Cestoni (1706) reported other parasitoids from eggs of cruciferous
insects.  He called aphids,
"cabbage 
     
sheep," and their parasitoids, "wolf mosquitoes."
 
G. 
Erasmus Darwin (1800) discussed the useful role of parasitoids and
predators in regulating insect pests.
 
IV.  During
the remainder of the 18th Century an ever increasing number of references to
entomophagous and 
      
entomogenous organisms appeared in the literature, largely in the form
of papers dealing with parasitoid biologies.
 
A. 
Diseases of silkworms were recognized early in the 18th Century.
 
B.  De
Reamur (1726) described and illustrated Cordyceps
fungus infecting a noctuid larva.
 
V.  Biological Control Efforts
in the 18th Century
 
A.  By
1762 the first successful importation of an organism from one country to
another for biological 
     
control took place with the introduction of the mynah bird from India to
the island of Mauritius, for locust 
     
control.  
 
B. 
Further development of modern biological control awaited the recognition
of the fact that insect pest 
     
problems were population phenomena. 
The controversial publications of Malthus appeared toward the 
      end
of the 18th Century, and generated considerable interest in the subject of
populations.  Malthus' work
      will be discussed further in the next section
on "Concepts in Population Ecology."
 
VI.  Biological Control Efforts
in the Early 19th Century
 
       A
number of articles appeared during the first half of the 19th Century that
lauded the beneficial effects 
of entomophagous insects.
 
A. 
Erasmus Darwin (1800) recommended protecting and encouraging syrphid
flies and ichneumonid 
     
wasps because they destroyed considerable numbers of cabbage-feeding
caterpillars.
 
B.  Kirby
& Spence  (1815) <PHOTO> showed that predaceous coccinellids
controlled aphids.
 
C.  Hartig
(1827) recommended the construction of large rearing cages for parasitized
caterpillars, with the 
     
ultimate aim of mass release.
 
D.  Ratzeberg
(ca. 1828) <PHOTO> called particular attention to the
value of parasitic insects with publication 
      of a
large volume on the parasitoids of forest insects in Germany.  He did not believe that parasitic control 
     
could be augmented by humans.
 
E. 
Agustino Bassi (1834) first demonstrated that a microorganism, Beauvaria bassiana, caused an animal 
     
disease, namely the muscardine disease of silkworms.
 
F. 
Kollär  (1837) <PHOTO> writing an article for farmers,
foresters and gardeners, pointed out the importance 
      of
entomophagous insects in nature's economy; studied parasitoid biologies and was
the first to report the
     
existence of egg parasitoids.
 
G. 
Boisgiraud (1843) reported that he used the predaceous carabid beetle, Calasoma sycophanta, to 
     
successfully control gypsy moth larvae on poplars growing near his home
in rural France.  He also reported
      that
he had destroyed earwigs in his garden by introducing predaceous staphylinid
beetles.
 
VII.  Biological Control in the Late 19th Century
 
      
Beginning in 1850, events associated with the westward expansion of agriculture
in the United States paved the 
way for the further development of the field of
biological control.  During and
following the "Gold Rush" in 
California, agriculture expanded tremendously in
California especially.  At first the new
and expanded plantings 
escaped the ravages of arthropod pests.  Predictably, however, crops soon began to
suffer from destructive 
arthropod outbreaks.  Many of these pests were found to be of foreign origin, and were
observed to be far more 
destructive in the newly colonized areas than in
their native countries.  Consequently,
the notion grew that perhaps
 these
pests had escaped from some regulatory factor or factors during their
accidental introduction into America.
 
A.  Asa
Fitch (1855) <PHOTO> was the State Entomologist of New York
who is recorded as the first entomologist 
      to
seriously consider the transfer of beneficial insects from one country to
another for the control of an 
     
agricultural pest.  Fitch suggested
that the European parasitoids of the wheat midge, Sitydiplosis mesellana,
be 
      sent
into the eastern United States.
 
B. 
Benjamin Walsh  <PHOTO> supported Fitch's suggestion and in
1866 he became the first worker in the United 
     
States to suggest that insects be employed in weed control.  He proposed that insects feeding on toad
flax, Linaria 
      vulgaris, be imported
from Europe to control invaded yellow toad flax plants.  The first actual case of biological 
     
control of weeds was, nevertheless, in Asia, where around 1865 the
cochineal insect Dactylopius ceylonicus was
      
introduced from southern India into Ceylon for prickly pear cactus
control (Opuntia vulgaris).  Originally Dactylopius 
      had been imported to India from Argentina in
1795, in the mistaken belief that it was the cochineal insect of commerce,
      D. cacti.
 
C.  Louis
Pasteur (1865-70) <PHOTO> studied silkworm diseases and saved the
silk industry in France from 
      
ruin [not really biological control].
 
D. 
Charles Valentine Riley (1870) <PHOTO>.
 
1.  The
father of modern biological control.
 
2.  He shipped
parasitoids of the plum curculio from Kirkwood, Missouri to other parts of that
state.
 
3.  In
1873 he became the first person to successfully transfer a predator from one
country to another with the
    
shipment of the American predatory mite, Tyroglyphus phylloxerae
to France for use against the destructive 
    
grapevine phylloxera.  The
results were not particularly successful, however.
 
4.  In
1883, Riley directed the first successful intercontinental transfer of an
insect parasitoid, Apanteles glomeratus,
     from
England to the United States for control of the imported cabbageworm.  He was Chief Entomologist of the 
     U. S.
Department of Agriculture.
 
5.  In
1872, 11 years before the importation of A.
glomeratus, Riley began his
interest in the cottony-cushion scale,
      Icerya purchasi, which was considered the most important citrus
pest in California.  He correctly
located its point 
     of
origin in Australia.
 
    
[Doutt's account of this biological control program on p. 31-38 of the
DeBach (1964) text is particularly colorful. 
      Read
this, paying particular attention to the following:
 
a.  the
roles played by Riley, Albert Koebele 
<PHOTO> and D. W. Coquillet <PHOTO>.
 
b.  note
the species of insects involved (the vedalia beetle, Rodolia cardinalis
<PHOTO>, and the dipterous 
    
parasitoid, Cryptochaetum
iceryae), their source,
numbers imported, and their activities relative to the cottony-
    
cushion scale.
 
c.  note
the method of colonization, and be able to describe the spectacular results of
these introductions, which 
   
changed the status of the pest to an insect of no economic importance in
only four years time.
 
VIII.  The
successful biological control effort against the cottony-cushion scale spirited
many biological control attempts
       in
many countries, resulting in over 200 biological control successes (see Chapter
24 of the DeBach (1964) text and other
      
hand-outs).
 
A.  The
cottony-cushion scale success admittedly harmed overall pest control in
California for quite some time 
     
because growers thought that the vedalia beetle would also control other
insect pests.  Consequently, they 
     
neglected other mechanical and chemical control methods.
 
B.  George
Compere (1899) became the first state employee specifically hired for
biological control work.  He
     
worked as a foreign collector until 1910, during which time he sent many
shipments of beneficial insects to
     
California from many parts of the world.   Harold Compere <PHOTO>, his son,
also devoted his entire career 
     to
the search for and identification of natural enemies of scale insects.
 
C.  Harry
Scott Smith (1913) <PHOTO>.
 
1.  He was
appointed superintendent of the State Insectary in Sacramento.
 
2.  In
1923, biological control work was transferred to the Citrus Experiment Station
and Graduate School of
     
Subtropical Agriculture of the University of California, Riverside.
 
3. 
Biological control work at Riverside was first conducted in the Division
of Beneficial Insect Investigations, 
     and
was changed to the Division of Biological Control with Smith as chairman in
1947.  Personnel were stationed 
     at
Albany and Riverside.
 
4. 
Importation of Chrysolina
beetles from Australia for Klamath weed control marked the beginning of
biological
     weed control
in California in 1944.
 
5.  Edward
Steinhaus (1947) <PHOTO> established the first laboratory and
curriculum in insect pathology at the
    
University of California, Berkeley. 
Later he transferred to the newly opened Irvine campus of the University
and
    
attempted to further insect pathology there.  His untimely death in 1968 precluded this goal.
 
6.  The
Division of Biological Control became the Department of Biological Control at
UC Riverside and Berkeley 
      in
1954.
 
7.  In
1969 Biological Control was dropped as a department, becoming a Division of
Biological Control within the
     
Department of Entomology, against the wishes of the entire biological
control faculty, numbering over 24 academics
      at Riverside and Berkeley at that
time.  
 
8.  The
Berkeley faculty created their own separate Division of Biological Control with
guaranteed privileges and
     
minimum control by the Department of Entomology.  At Riverside the Division of Biological
Control gradually 
    
became dominated by chemical control oriented faculty in the Department
of Entomology.  In 1989 the Division 
     was
abolished, against the wishes of 85% of the faculty in the Division.  Ignorance and pecuniary control among 
     the ranks of University of California
bureaucrats is believed to be the principal cause.  Although the dissenting 
   
faculty in the Division each wrote a personal plea to the then
Chancellor Rosemary S. J. Schraer to discuss the 
   
matter, in not one case was a reply received.  
 
 
REFERENCES:
 
Bassi, A.  1935. 
Del mal del segno, calcinaccio o moscardino, mallatia che affigge i
bachi da seta e sul modo di liberarne le bigattaie anche le piu infestate.  Part I: 
Theoria.  Orcesi, Lodi. p. 1-9,
1-67.
 
Bodenheimer, F. S.  1931. 
Der Massenwechsel in der Tierwelt. 
Grundriss einer allgemeinen tierischen Bevölkerungslehre.  Arch. Zool. Ital. (Napoli) 16:  98-111.
 
Compere, G.  1902. 
Entomologist's Report. 
Introduction of Parasites.  West.
Austral. Dept. Agric. J. 6:  237-40.
 
Compere, G.  1904. 
Black scale parasite (Scutellista
cyanea).  West Austral. Dept. Agric. J. 10:  94.
 
Compere, G.  1921. 
Seasonal history of black scale and relation to biological control.  Calif. Citrog. 6:  197.
 
Darwin, E.  1800. 
Phytologia.  Publ., London.
 
DeBach, P.  1974. 
Biological control by natural enemies. 
Combridge Univ. Press.
 
Doutt, R. L.  1964. 
The historical Development of biological control.  In:  P. DeBach (ed.), Biological Control of
Insect Pests and Weeds.  Reinhold Publ.
Corp., New York.  844 p.
 
Fitch, Asa.  1954. 
Sixth, seventh, eighth and ninth reports on the noxious, beneficial and
other insects of the state of New York. 
Albany, New York.  259 p.
 
Goedaert, J.  1662. 
Metamorphosis et Historia Naturalis Insectorum.  Jacques Fierens, Middelburgh.  
 
Kirby, W. & W.
Spence.  1815.  An Introduction to Entomology. 
Longman, Brown, Green & Longmans, London.  285 p.
 
Kollär, Vincent.  1837. 
In:  London's Gardner's Magazine. 1840.  [English translation].
 
Malthus, T. R.  1803. 
An Essay on the Principle of Population as It Affects the Future
Improvement of Society.  J. Johnson,
London, 2nd ed.  610 p.
 
Pasteur, L.  1870 
Etudes dur la maladie des vers a soie. 
Gautherie-Villars, Paris, I:  322
p.; II: 327 p.
 
Ratzeburg, J. T. C.  1944a. 
Die Ichneumonen der Forstinsekten in forstlicher und entomologischer
Beziehung; ein Anhang zur Abbildung und Beschreibung der Forstinsekten.  Theile, Berlin.  3 vol.
 
Ratzeburg, J. T. C.  1944b. 
Die Ichneumonen der Forstinsekten, Vol. I.  Berlin.
 
Réaumur, M. de.  1726. 
Remarques sur la plante appellée a la Chine Hia Tsao Tom Tchom, ou
plante ver.  Mem. Acad. Roy. Sci. (21
Aug 1726).  p. 302-5.
 
Riley, C. V.  1893. 
Parasitic and predaceous insects in applied entomology.  Insect Life 6:  130-41.
 
Riley, W. A.  1931. 
Erasmus Darwin and the biologic control of insects.  Science 73: 
475-6.
 
Smith, H. S.  1916. 
An attempt to redefine the host relationships exhibited by entomophagous
insects.  J. Econ. Ent. 9:  477-86.
 
Smith, H. S.  1919. 
On some phases of insect control by the biological method.  J. Econ. Ent. 12:  288-92.
 
Smith, H. S.  1929. 
The utilization of entomophagous insects in the control of citrus
pests.  Trans. 4th Internatl. Congr.
Ent. 2:  191-8.
 
Steinhaus, E. A.  1946. 
Insect Microbiology.  Comstock
Publ. Co., Inc., Ithaca, New York.  763
p.
 
Steinhaus, E. A.  1949. 
Principles of Insect Pathology. 
McGraw-Hill Book Co., Inc., New York. 
757 p.
 
van Leeuwenhoek, A.  1702. 
Letter in Nr. 266 of the Philosophical Transaction 1700-1701, Vol. 22,
p. 659-72.  Smith & Walford, London.
 
van Lenteren, J. C.  1983. 
Biological pest control:  passing
fashion or here to stay?  Organorama
(Netherlands) 20:  1-9.
 
Walsh, B. D.  1866. 
Practical Entomologist.  June
1866.  p. 101.