H
Habitat
The place where an organism dwells. This is a less
inclusive term than “niche.”
Habitat Diversity
The range of habitats present in an area. Because
insects often have preferred habitats, there usually
is a strong correlation between habitat and insect
diversity.
Habituation
Failure to elicit normal response after repeated
stimuli. Often the organism gradually decreases it
response to the stimulus, and the organism may
not respond even after the stimulus is discontin-
ued for a protracted period. All insects are thought
to be capable of habituation to some stimuli, and it
is considered to be the simplest form of learning.
 Associative Learning
 Latent Learning
 Insight Learning
 Learning in Insects
Haddow, Alexander John
Alex Haddow was born in Scotland in December
1912, and as a boy showed intense interest in insects.
His first degree was in zoology, from Glasgow
­ niversity in 1934. He took another 4 years to com-
U
plete an M.D. degree. Next, he studied at the London
School of Tropical Medicine for a diploma in tropi-
cal medicine. In 1941 he traveled to Kenya and stud-
ied biting cycles of mosquitoes under conditions of
a 24-h continuous catch, developing and standard-
izing methods for collecting and analyzing the data.
Other work in Africa was on the epidemiology of
yellow fever, and it involved collecting and sampling
from monkeys. After 24 years in Africa, he returned
to Britain in 1965, to become Administrative Dean
of the Faculty of Medicine at Glasgow University.
His honors include election to the fellowship of
The  Royal Society as well as the Royal Society of
Edinburgh. He died in Glasgow on December 26,
1978, survived by his wife, Peggy, and two sons.
Reference
Gillett JD (1979) Alexander John Haddow 1912–1978.
Antenna 3:54
Haematomyzidae
A family of chewing lice (order Phthiraptera).
 Chewing and Sucking Lice
Haematopinidae
A family of sucking lice (order Phthiraptera). They
sometimes are called ungulate lice.
 Chewing and Sucking Lice
1762
H Hagen, Hermann August

Hagen, Hermann August

Hermann Hagen (Fig. 1) was born in Königsberg
(now Kaliningrad), then in German East Prussia
(now in Russia), on May 30, 1817. He was educated
at Universität Königsberg, and became a physician
in  that city. He studied extant insects (Odonata,
Neuroptera, Isoptera, Psocoptera, Plecoptera,
and Trichoptera) and fossil insects. Some of his
major works were “Monographie der Termiten”
(1855–1860) and “Bibliotheca Entomologica”
(1862–1863) [for a subsequent bibliography of
the  world’s entomological literature, see: Horn,
Walther]. In 1867 he was invited to the USA to
take charge of the entomological section of the
Museum of Comparative Zoology at Harvard
University in Massachusetts. He accepted, orga- Hagen, Hermann August, Figure 1  Hermann
nized and built that collection, and greatly influ- Hagen.
enced taxonomic entomologists in the USA. In
1890–1891 he was afflicted with paralysis and entomologist in the Division of Biological Con-
influenza, and died in Massachusetts in 1893. trol, of the University of California’s Agricultural
Experiment Station at Albany. He became ento-
mologist in 1965 and professor of entomology in
References 1969. His areas of research interest included the
behavior of coccinellids, highlighted in a (1970)
Essig EO (1931) Hagen, Hermann August. In: A history of article in National Geographic called “The high-
entomology. Macmillan, New York, NY, pp 643–646
Mallis A (1971) Hermann August Hagen. In: American ento-
flying ladybug.” He was the leader of projects
mologists. Rutgers University Press, New Brunswick, NJ, that resulted in introduction of 25 species of bio-
pp 119–126 control agents against 17 economically impor-
tant pests. He had a pioneering role in the
nutrition of biocontrol agents, with proof in
the field that manipulative techniques using arti-
Hagen, Kenneth Sverre ficial nutrition could work. He authored more
than 160 scientific publications and received
Ken Hagen was born in the state of California, numerous awards. He was also a gifted teacher.
USA, on November 26, 1919. He displayed an He died on January 10, 1997, survived by his
early interest in insects and formed a collection. wife, Maxine, and one son.
He earned a B.S. degree from the University of
California at Berkeley in 1943. After service
in  the U.S. navy in World War II (landings at
Normandy and Okinawa), he returned to the References
graduate school of the same institution, earning
Caltagirone LE, Dahlsten DL, Garcia R (1997) Kenneth Sverre
degrees of M.S. in 1948 and Ph.D. in 1952. While
Hagen, Entomological sciences: Berkeley. Available at
he was a graduate student he worked as a techni- http://sunsite.berkeley.edu:2020/dynaweb/teiproj/uchist/
cian, but after his Ph.D. he was hired as junior inmemoriam/inmemoriam1997. Accessed Aug 2002

Hahn, Carl Wilhelm
Carl Hahn was born on December 16, 1786, and
lived mainly in Nürnberg (Nuremberg), Germany.
He studied the spiders of his country, which he
described and illustrated in two major works,
“Monographie der Spinnen” and “Die Arachniden”.
His premature death in 1836 interrupted publica-
tion of the latter, of which he completed only two
volumes. It was then completed, in a further 14
volumes, by C. L. Koch between 1836 and 1849.
Reference
Bonnet P (1945) Bibliographia Araneorum 1:32
Hair Pencil
Clusters of long setae on the body of certain insect
(particularly male Lepidoptera and some Neu-
roptera). They occur on various parts of the body,
but usually on the abdomen. They are associated
with exocrine glands, and usually used during
courtship to disperse sex pheromones. This term is
synonymous with “brush organ.”
Hair Plate
A sensor structure found at the leg joints and at
other limb articulations. They respond to touch,
bending and joint flexing by emitting nervous
stimuli, and adapt slowly. They allow the insect to
know the orientation of the head and appendages.
A more complex version of the hair plate is the
chordotonal sensillum.
Hairstreaks
Some members of the family Lycaenidae (order
Lepidoptera).
 Gossamer Winged Butterflies
 Butterflies and Moths
Hale Carpenter, Geoffrey Douglas
H 1763
Hairy Chinch Bug, Blissus
leucopterus hirtus Montandon
(Hempitera: Lygaeidae)
Hairy chinch bug is an increasingly important pest
of turfgrass.
 Turfgrass Insects and their Management
Hairy Fungus Beetles
Members of the family Mycetophagidae (order
Coleoptera).
 Beetles
Hale Carpenter, Geoffrey Douglas
Geoffrey Hale Carpenter was born in Eton Col-
lege, England, on October 26, 1882. His under-
graduate studies were at Oxford University, and
then in 1908 he qualified in medicine and took a
D.M. degree in 1913. But by 1911, interested in
tropical medicine and natural history, he was
studying the bionomics of Glossina palpalis and
its relation to sleeping sickness in Uganda. His
research was published in reports of The Royal
Society in 1912, 1913, and 1919. In Uganda, too,
he studied mimicry in butterflies, and through
rearing experiments solved problems of relation-
ships. He viewed mimicry among butterflies as a
result of natural selection by predatory birds,
mammals, and reptiles. His butterfly studies are
published mainly in journals of the Royal Ento-
mological Society. In 1933 he was appointed Hope
Professor at Oxford University, succeeding Sir
Edward Poulton. He retired in 1948 and died in
Oxford on January 30, 1953.
Reference
Riley ND, Hale Carpenter GD (1953) Entomologist
86:155–156
1764
H Half Life
Half Life
The period of time required for a pesticide to lose
half of its original effectiveness or toxicity.
Halictidae
A family of bees (order Hymenoptera, superfamily
Apoidae).
 Wasps, Ants, Bees and Sawflies
Halictophagidae
A family of insects in the order Strepsiptera.
 Stylopids
Haliday, Alexander Henry
Alexander Haliday (Fig. 2) was born in Belfast in
1807. At the age of 15, he entered Trinity College,
­Dublin, remaining there 5 years and obtaining an
M.A. degree. Next, he studied law, and apparently
was successful, although it is not clear that he ever
practiced this profession. Returning to the north of
Ireland, he devoted his time to the study of litera-
ture and natural history. He was appointed High
Sheriff of the county of Antrim in 1843, a political
Haliday, Alexander Henry, Figure 2  Alexander
Haliday.
position rather than one in law-enforcement.
Beginning in 1828, he published a long series of
papers on the Irish insect fauna, especially Diptera.
He also published extensively on Chalcidoidea
and other “parasitic” Hymenoptera, and on Thys-
anoptera. Due to poor health, he emigrated to
Lucca, Italy, about 1860. There, he continued
collecting insects and helped to found the Società
Entomologica Italiana. He died on July 12, 1870.
Reference
Anonymous (1870) Alexander Henry Haliday. Entomologist’s
Monthly Magazine 7:91
Halimococcidae
A family of insects in the superfamily Coccoidae
(order Hemiptera).
 Bugs
Haliplidae
A family of beetles (order Coleoptera). They com-
monly are known as crawling water beetles. Beetles
Haltere
(pl., halteres) The vestigial, modified hind wings of
Diptera that function as balancing organs.
Hamophthiriidae
A family of sucking lice (order Phthiraptera).
 Chewing and Sucking Lice
Hamulus
(pl., hamuli) Hook-shaped hairs on the leading
edge of the hind wing in Hymenoptera that unite
the fore- and hind wings.
 Wings of Insects
 Haploid
H 1765

Handling Time Hangingflies

The time a predator spends pursuing, subduing, Members of the family Bittacidae (order
and consuming prey (a meal). Mecoptera).
 Scorpionflies

Handlirsch, Anton
Hansen, Viktor
Anton Handlirsch was born in Vienna on January
Viktor Hansen was born in Copenhagen on August
20, 1865. He studied pharmacy, and in 1883
29, 1889. In 1907 he entered the Metropolitanskolen
received a master’s degree in that subject. Then,
and in 1913 obtained his law degree. He worked in
he became scientific helper, assistant (1892),
the Justitsministeriet from 1915, became a superior
adjunct custodian (1899), second-class custo-
judge in 1941, and worked in that capacity until he
dian (1906), first-class custodian (1918) and
retired in 1959. However, he began to collect beetles
director (1922) of the Naturhistorisches Hofmu-
as a teenager, and in 1905 joined the Entomologisk
seum of Vienna. His earliest entomological work
Forening [entomological society]. His first paper
was on Hymenoptera, reaching a climax (1887–
was published in 1907 in Entomologiske Mededelser.
1894) with a monograph of wasps related to
His lifetime total was more than 100 papers on the
Bembex and Nysson. He took charge of Hemiptera
Danish beetle fauna. His major production was 23
at the museum, purchased the Signoret collec-
volumes on beetles in the series Danmarks Fauna.
tion for it, and wrote a monograph on Phymati-
For this, he was awarded an honorary doctoral
dae (1897). Next, he turned his attention to fossil
degree from Københavns Universitet in 1950, and
insects. His (1906–1908) book “Die fossilen
medals from entomological societies in Denmark
Insekten und die Phylogenie der rezenten For-
and Sweden, and from a natural history society in
men” enumerated or described all known fossil
Denmark. He died on March 6, 1974.
insects. It included a new classification with dis-
cussion. In his contributions to Schroeder’s (1925)
“Handbuch der Entomologie” and Kükenthal’s Reference
(1926–1935) “Handbuch der Zoologie,” he mod-
ified the classification. He also published about Herman LH (2001) Hansen, Viktor. Bull Am Mus Nat Hist
100 other titles. He died in Vienna on August 265:75
28, 1935.
Haplodiploidy
Reference A type of parthenogenetic reproduction in which
males are produced from unfertilized eggs and
Calvert PP (1936) Obituary. Entomological News
47:168–169
are  haploid, while the females are diploid. In
Hymenoptera, females can control the release of
sperm, and regulate the sex ratio of offspring.
Handsome Fungus Beetles
Haploid
Members of the family Endomychidae (order
Coleoptera). Cells or organisms that contain a single copy of
 Beetles each chromosome.
1766
H Hardy-Weinberg Equilibrium
Hardy-Weinberg Equilibrium
An equilibrium of genotypes achieved in popula-
tions of infinite size in which there is no migra-
tion, selection, or mutation after at least one
generation of panmictic mating. With two alleles,
A and a, of frequency p and q, the Hardy-Weinberg
equilibrium frequencies of the genotypes AA, Aa
and aa are p2, 2pq and q2, respectively.
Harlequin Bug
 Crucifer Pests and their management
Harlequin Bug, Murgantia
histrionica (Hahn) (Hemiptera:
Pentatomidae)
john l. capinera
University of Florida, Gainesville, FL, USA
Indigenous to Mexico and Central America,
­harlequin bug has dispersed north into the United
States. Its appearance in Texas, USA, in 1864
­coincided with the occurrence of Union troops
during the American Civil War, and in parts of
the South earned it the name “Sherman-bug” after
the northern General Sherman, and “Lincolnite”
after President Abraham Lincoln. It rapidly
spread throughout the southern states, and even-
tually reached northern locales such as Colorado,
Iowa, southern Michigan, Pennsylvania, and
­Massachusetts. It is considered to be a serious pest
only in southern states, however, and is not
regarded as a problem in California. It has also
dispersed to the Hawaiian Islands.
Life History
Harlequin bug breeds continuously in the south-
ern portions of its range. During mild winters all
stages have been observed as far north as Virginia.
In colder climates only the adults survive the win-
ter in sheltered locations. They seek shelter in and
near fields, among overwintering crop plants,
and  in other organic debris such as dead leaves
and bunches of grass. Two or three generations
per year seem to be normal, but there appears to
be four generations in south Texas.
Adults begin depositing eggs about 2 weeks
after becoming active in the spring. Eggs are depos-
ited beneath leaves, usually in clusters of 12 arranged
in two rows of six, at intervals of 5–6 days. As the
female nears the end of her life, the egg batches get
slightly smaller and the egg arrangement less regu-
lar. The eggs are barrel-shaped, and measure about
1.30–1.38 mm long and 0.90–0.92 mm in diameter.
They are light gray or pale yellow in color, and gen-
erally are circled by two black bands. They may also
bear small black dots or spots, and the top has a
semicircular black marking. The average number of
eggs is reported to be 115 per female. Egg deposi-
tion may occur over a period of 40–80 days. Eggs
hatch in about 4–5 days during warm weather, while
15–20 days may be required during cool weather.
Upon hatching, young nymphs stay clustered
near the old eggs for 1 or 2 days. The newly hatched
nymphs are pale green with black markings, but
soon become brightly colored: black or blue, with
red and yellow or orange markings. Reportedly
there are six instars in Texas, and nymphal develop-
ment can be completed in as little as 30 days. There,
average development times are 3.4, 3.2, 4.7, 4.7, 7.0,
and 4.3 days, respectively, for the six instars devel-
oping under summer conditions. Under spring
conditions, development times were in­­creased by
about 30%. Studies in Virginia suggest only five
instars, however, and a development time require-
ment of 40–60 days during the summer, and slightly
longer, perhaps 70 days, during cool weather.
The adults usually live about 60 days, but may
live considerably longer during the winter. They
measure about 8.0–11.5 mm in length. The adults
are brightly colored, similar to the large nymphs,
principally black and yellow or black and red. The
color pattern varies (Figs. 3 and 4), with the spring
and summer bugs being more brightly colored
 Harlequin Bug, Murgantia histrionica (Hahn) (Hemiptera: Pentatomidae)
Harlequin Bug, Murgantia histrionica (Hahn)
(Hemiptera: Pentatomidae), Figure 3  Adult of
harlequin bug, Murgantia histrionica (Hahn).
Harlequin Bug, Murgantia histrionica (Hahn)
(Hemiptera: Pentatomidae), Figure 4  Third instar
of harlequin bug, Murgantia histrionica (Hahn).
than the overwintering insects. As with many stink
bugs, harlequin bugs produce a disagreeable odor
if disturbed, and birds avoid eating them.
Harlequin bug is principally a pest of crucifers,
attacking broccoli, Brussels sprout, cabbage,
cauliflower, Chinese cabbage, collard, kale, kohlrabi,
mustard, radish, rutabaga, turnip, and watercress.
Harlequin bug is reported to be especially fond of
H 1767
horseradish. In the southernmost states, crucifers do
not thrive during the summer months, and the bugs
are forced onto other plants. Thus, they are sometimes
found feeding on asparagus, beans, okra, squash,
tomato and many other vegetables, but this is usu-
ally due to lack of normal food. ­Harlequin bug feeds
readily on cruciferous weeds such as wild mustard,
Brassica spp.; shepherds purse, Capsella bursa-pasto-
ris; and pepperweed, Lepidium spp.; and related
mustard oil-containing plants such as members of
the family Capparaceae. Other weeds common in
crops, such as pigweed, Amaranthus spp., and lamb-
squarter, Chenopodium album, are also fed upon,
and reproduction occurs on these plants.
Harlequin bug appears to be relatively free of
natural enemies, other than for egg parasites and
general predators. The egg parasitoids are Oen-
cyryus johnsoni (Howard) (Hymenoptera: Encyrti-
dae), Trissolcus murgantiae Ashmead, and T. podisi
Ashmead (both Hymenoptera: Scelionidae). The
species that is best known is O. johnsoni, which has
been reported frequently from harlequin bug eggs,
and has caused up to 50% mortality during a har-
lequin bug outbreak in Virginia. This parasite is
widely distributed, and apparently has other hosts.
It attacks eggs in all stages of embryonic develop-
ment, and prevents the eggs from hatching. How-
ever, O. johnsoni is not the only effective parasite,
as T. murgantiae was observed to parasitize 45% of
harlequin bug eggs in North Carolina, at locations
where O. johnsoni parasitized only 30% of eggs.
Because of its effectiveness, T. murgantiae was
introduced into California.
Damage
The piercing-sucking feeding behavior of this insect
results in white blotches at the site of feeding. Wilt-
ing, deformity, and plant death may occur if insects
are abundant. Mild winters are said to favor survival,
and subsequent damage. Once considered the most
serious crucifer pest in the south, this insect has
been relegated to minor status in commercial pro-
duction and persists mostly as a home-garden pest.
1768
H Harris, Thaddeus William

Management medicine and married. In 1823 he published his

Insecticides are applied to the foliage for suppres-
sion of harlequin bug. Harlequin bug can be diffi-
cult to control with insecticides; targeting the
young bugs and thorough coverage are recom-
mended. Soap applied alone or in combination
with rotenone has provided good control.
Trap crops, usually consisting of early-planted
mustard, rape, or kale are sometimes recom-
mended to divert the overwintering bugs from the
principal crop. Such trap crops must be sprayed or
destroyed, however, or the adults will soon move
to the main crop. Destruction of crop residues, on
which the insect may overwinter in the north or
oversummer in the south, is an important cultural
practice to alleviate harlequin bug damage.
Susceptibility to damage varies among cruci-
fer crops. Mustard and Chinese cabbage are quite
susceptible; turnip, kale, rutabaga, and some rad-
ishes are intermediate; and cauliflower, cabbage,
broccoli, collard, Brussels sprout, kohlrabi, and
most radish varieties are fairly resistant. Cabbages
are the most resistant crop, but considerable varia-
tion among cultivars is evident.
 Crucifer Pests and their Management
 Vegetable Pests and their Management, Stink
Bugs (Hemiptera: Pentatomidae) Emphasizing
Economic Importance
first entomological paper. In 1831 he published a
catalogue of insects and became librarian at Har-
vard University. Teaching of courses in natural
history and entomology followed. His 1841 “Report
on insects injurious to vegetation” became a classic
and was reprinted several times. He also published
some descriptions of new species of insects
that  were of economic importance. He died in
Massachusetts on January 16, 1856.
References
Essig EO (1931) Harris, Thaddeus William. In: A history of
entomology. Macmillan, New York, NY, pp 651–653
Mallis A (1971) Thaddeus William Harris. In: American ento-
mologists. Rutgers University Press, New Brunswick, NJ,
pp 25–33
Harvester Ants
Ant species that store seeds in their nests. This
behavior occurs in many ant taxa, including some
that are not closely related.
 Harvester Ants, Pogonomyrmex

References

Capinera JL (2001) Handbook of vegetable pests. Academic

Press, San Diego, 729 pp
White WH, Brannon LW (1939) The harlequin bug and its
control. USDA Farmer’s Bulletin 1712, 10 pp

Harris, Thaddeus William

Thaddeus Harris (Fig. 5) was born in Massachu-

setts on November 12, 1795. He obtained a first
degree from Harvard University in 1815 and then Harris, Thaddeus William, Figure 5  Thaddeus
a medical degree in 1820, after which he practiced Harris.
 Harvester Ants, Pogonomyrmex Mayr (Hymenoptera: Formicidae)
Harvester Ants, Pogonomyrmex
Mayr (Hymenoptera: Formicidae)
thomas o. crist
Miami University, Oxford, OH, USA
Harvester ants are so named because they collect
and store seeds in their nests for later consump-
tion. The seed-harvesting habits of ants are noted
in several historical accounts and are the subject of
numerous scientific studies. Over 150 species of
seed-harvesting ants occur worldwide. In the New
World, harvester ants comprise 60 species belong-
ing to two closely related genera, Pogonomyrmex
Mayr (“bearded ant”) and Ephebomyrmex Wheeler
(“youthful ant”). The beard, or psammophore,
refers to the tuft of hairs that extend below the head
on the workers of nearly all 45 species of Pogono-
myrmex. This distinctive trait corresponds to the
ground-nesting habits of harvester ants because
the psammophore is used to move particles of soil.
Some harvester ants build large mounds or craters
of soil and gravel on the soil surface while others
have more inconspicuous nests under rocks.
Most Pogonomyrmex occur in deserts and
grasslands where seeds are abundant, but a few
species are found in forest and montane environ-
ments. Harvester ants (Fig.  6) range throughout
South America, portions of Central America and
the Caribbean, and virtually all of western North
America and the southeastern Coastal Plain. The
geographic ranges of most species, however, are
found within southwestern North America and
northern South America, the latter of which is
likely the evolutionary origin of Pogonomyrmex.
The mating behavior and life cycle is generally
similar among Pogonomyrmex species. During
spring or summer, ant colonies produce numerous
winged males and females (“alates”) that are repro-
ductively viable and larger than the sterile workers.
Alate flight activity is synchronous among colonies,
usually occurring in the late morning on 2 or 3 days
following a significant rainfall. Mating flights can
occur several times a year, but are ­usually restricted
to a 2 or 3-week period. After leaving the nest, alates
H 1769
Harvester Ants, Pogonomyrmex Mayr
­(Hymenoptera: Formicidae), Figure 6  A worker
harvester ant. The genus Pogonomyrmex is named
after the prominent beard (the psammophore)
on the lower side of the head. Drawing is by
Ruth Ann DeNicola, published in Cole, A. C., Jr.
(1968) Pogonomyrmex harvester ants: a study of
the genus in North America. Reproduced with
­permission from The University of Tennessee
Press, Knoxville, Tennessee.
aggregate on hills or cliffs and in the uppermost por-
tions of shrubs, trees, or structures such as fence
posts, windmills, or buildings. Males often “lek”
(form mating aggregations) together in groups of
5–20 individuals, and females are attracted to male
aggregations by a pheromone. Females typically
mate with several males, a behavior that results
in genetically variable workers after the queen begins
reproduction. In some species, such as P. badius (the
Florida harvester ant), lekking does not occur and
males fly to another nest to mate with females, or
mating might take place between males and females
of the same colony, which can result in inbreeding
and a loss of genetic variability over time.
After mating, the fertilized female sheds her
wings and begins excavating a nest in the soil. Most
“foundress queens” die before successfully estab-
lishing a colony. The queen must carry out all of the
worker tasks – foraging, nest maintenance, and
brood care – for 2 or 3 weeks until her first brood
matures into adult workers. The mortality of young
colonies continues to be high until the next year
when the size of the worker population may increase
substantially. Pogonomyrmex colonies have variable
1770
H Harvester Ants, Pogonomyrmex Mayr (Hymenoptera: Formicidae)
numbers of workers: P. laticeps and P. magnacanthus
have only 50–250, whereas P. rugosus (the rough
harvester ant) and P. barbatus (the red harvester
ant) may boast up to 15,000. Most species have a
worker force of 500–5,000 ants. Two species, the
slavemaking P. anergismus and P. colei, have no
workers at all; instead, they live in the nests of
P. rugosus or P. barbatus and enslave workers to raise
reproductive males and females for them. It appears
that all Pogonomyrmex species are monogynous
(have one queen) during the life of a colony, which
may persist for 10–50 years in some species such
as  P. occidentalis (the western harvester ant) and
P. owyheei [salinus] (the Owyhee harvester ant).
Workers perform a variety of tasks. Nest work-
ers take care of the queen and developing larvae
and pupae. They also create and maintain a variety
of nest chambers for seed storage, brood develop-
ment and discarded refuse. Storage chambers are
connected by networks of tunnels, which may
extend 2–3 m vertically below ground in some spe-
cies. The queen and workers usually overwinter
more than a meter below ground in colder climates.
Worker tasks vary according to the age of the worker
and the needs of the colony. After emerging from
pupae, ants work mostly within the nest. Then, as
workers get older, they shift to exterior maintenance
of the nest, and finally to patrolling, scouting, or for-
aging, which may entail venturing up to 20 m from
the nest in some species. Workers also switch tasks
according to the needs of the colony. For example, if
a nest is damaged by heavy rain, foraging workers
might switch to nest maintenance. Similarly, a nest
intruder may elicit an alarm pheromone that rap-
idly employs a substantial number of workers to
defend the nest. Harvester ants are well-equipped
for nest defense, as the workers of many species will
deliver a ­painful sting.
A number of scientific studies have examined
the foraging and seed-harvesting behavior of Pogon-
omyrmex ants. Foraging workers search for seeds in
an individualistic manner, specializing on particular
seed species or locations near the nest. Although this
is the primary foraging strategy in   some species
(e.g., P. desertorum, P. maricopa, and P. californicus,
the California harvester ant), several harvester ants
also show group foraging tactics. These involve the
use of permanent pathways (trunk trails) that orient
workers to different foraging areas, and pheromone
trails that chemically recruit workers to areas of
high-food density (e.g., P. rugosus, P. barbatus, and
P. occidentalis). Recruitment pheromones are volatile
compounds that dissipate rapidly if workers do not
reinforce them by laying additional pheromones
along the recruitment trail. Ants usually harvest
<10% of the total seeds available in their foraging
areas, but they often remove a much larger fraction
of the seed species they prefer – grasses and small-
seeded forbs – and consequently, influence the types
of plants that are found near nests. A significant part
of the harvester ant diet is arthropods, especially
in some species found at higher elevations (e.g.,
P. montanus and P. mayri). A variety of other materials
are also returned to the nest by workers, including
twigs, leaves, flowers and feces.
Several vertebrates and arthropods prey on
harvester ants. Most commonly, these are birds, liz-
ards and spiders. Horned lizards (Phrynosoma spp.)
are specialist predators of harvester ants and have
evolved a resistance to the toxic compounds in the
venom delivered by the sting of worker ants. Preda-
tion by horned lizards can substantially reduce the
number of workers in a nest. Ants respond to lizard
predation by reducing their foraging activity, or
closing their nests entirely. The storage of seeds in
harvester ant nests is thought to provide a food
source when foraging activity is decreased in
response to predation. Other seed-eating animals,
such as kangaroo rats (Dipodomys spp.), may also
compete with harvester ants for food.
The large nest size and colony longevity of
many Pogonomyrmex species have important
­consequences to soils and plants. Ants change
the  physical and chemical characteristics of soil
by  increasing the porosity, organic matter and
nutrient levels. The alteration of nest soils and the
removal of plants and seeds by ants result in differ-
ent plant species near nests compared to surround-
ing areas. Therefore, in deserts and grasslands
where colony densities are high, harvester ants

influence plant species diversity and composition.
Recently, however, the native Pogonomyrmex ants
have been displaced in several areas by more
aggressive, invasive species such as the red imported
fire ant (Solenopsis invicta), or the Argentine ant
(Linepithema humile). A loss of harvester ants from
these ecosystems may have several consequences
to the constituent soils, plants and animals.
The biology and behavior of Pogonomyrmex
ants continues to fascinate scientists and amateurs
alike. Most studies of harvester ants are from rela-
tively few of the 60 species; future investigations of
lesser-known species will further our understand-
ing of these remarkable insects.
 Ants
References
Cole AC Jr, (1968) Pogonomyrmex harvester ants: a study of
the genus in North America. University of Tennessee
Press, Knoxville, Tennessee
Gordon DM (1995). The development of organization in an
ant colony. Am Sci 83:50–57
Hölldobler B, Wilson EO (1990) The ants. The Belknap Press
of Harvard University Press, Cambridge, MA
MacKay WP (1990) The biology and economic impact of
Pogonomyrmex ants. In: Vander Meer RK, Jaffe K,
Cedeno A (eds) Applied myrmecology: a world perspec-
tive, Westview Press, Boulder, Colorado, pp 533–542
MacMahon JA, Mull JF, Crist TO (2000) Harvester ants
(Pogonomyrmex spp.): their community and ecosystem
influences. Ann Rev Ecol Syst 31:265–291
Taber SW (1998) The world of the harvester ants. Texas A&M
University Press, College Station, Texas
Harvesters
Some members of the family Lycaenidae (order
Lepidoptera).
 Gossamer-Winged Butterflies
 Butterflies and Moths
Hatch, Melville Harrison
Melville Hatch was born in Detroit, Michigan,
USA,  on November 25, 1898. An early interest in
Hawk Lice
H 1771
biology led to his seeking and obtaining a university
education in the subject, culminating in a Ph.D.
from the University of Michigan in 1925. He became
a teaching assistant at the University of Michigan
in  1926, then an instructor at the University of
­Minnesota, and next an assistant professor at the
University of Washington. Promotions followed, and
he became professor in 1941 and chairman of the
Zoology Department, and Curator of Entomology
at the university’s Burke Museum in 1962. He retired
from teaching in 1969. Meanwhile he had begun
to write and publish his greatest work, “The beetles
of the Pacific Northwest,” which appeared in five
volumes, the last in 1971. His other publications
numbered more than 170, and he served as editor of
“The ­Biologist” from 1959 through 1967. His
Coleoptera collection was moved to the University
of Oregon after he had ceased to work on it. He died
near ­Seattle, Washington, on January 19, 1988.
References
Becker EC (1993) Melville H. Hatch (1898–1988). Coleopter-
ists Bull 147:112
Herman LH (2001) Hatch, Melville Harrison. Bull Am Mus
Nat Hist 265:75–76
Haustellate
This term is used to refer to mouthparts that are
formed for sucking, or piercing and sucking.
 Mouthparts of Hexapods
Haustellum
The portion of the mouth through which liquid
food is imbibed.
 Mouthparts of Hexapods
Hawk Lice
Members of the family Laemobothriide (order
Phthiraptera).
 Chewing and Sucking Lice
1772
H Hawk Moths (Lepidoptera: Sphingidae)
Hawk Moths (Lepidoptera:
Sphingidae)
John B. Heppner
Florida State Collection of Anthropods,
­Gainesville, FL, USA
Hawk moths, family Sphingidae (also called sphinx
moths, bee moths and hummingbird moths), total
1,230 species worldwide. Tropical regions of the
New World, Africa and Asia have the most biodi-
versity. There are three subfamilies: Smerinthinae,
Sphinginae, and Macroglossinae (sometimes only
two subfamilies are used). The family comprises
the monobasic superfamily Sphingoidea, in the
section Cossina, subsection Bombycina, of the
division Ditrysia. Adults (Fig.  7) medium size to
very large (23–200 mm wingspan), with head ver-
tex scaling mostly normal but sometimes rough-
ened; haustellum usually very long, to 30 cm (rarely
vestigial); labial palpi mostly upcurved (sometimes
porrect), with small third segment; maxillary palpi
small; antennae mostly clavate or lamellate and
thickened; body very robust (sometimes with lon-
ger hair-like setae); some genera with long needle-
like spines on hind tibiae (e.g., Oxyambulyx species
from Southeast Asia). Wings elongated and usually
with acute apex; hindwings usually elongated but
basally rounded, and much smaller than forewings.
Maculation varied but many with shades of brown
and gray, often with few markings, but also very
Hawk Moths (Lepidoptera: Sphingidae),
­Figure 7  Example of hawk moths (Sphingidae),
Callambulyx rubricosa (Walker) from Indonesia.
colorful species; some with hyaline wings and
mimicking wasps. Adults nocturnal or crepuscular
but some a diurnal. Larvae are leaf feeders, usually
with a posterior tail-like scolus; many larvae
extremely large. Host plants recorded in numerous
plant families. A few are economic. Among the larg-
est species in the family are females of ­Clanis titan
Rothschild & Jordan, of Southeast Asia, and Coe-
quosa triangularis Donovan, of ­Australia, both also
with massive bodies, while the smallest is Sphin-
gonaepiopsis obscurus Mabille, from Madagascar.
References
D’Abrera B (1986) Sphingidae mundi: hawk moths of the
world. E. W. Classey, Farringdon, 226 pp
Hodges RW (1971) Sphingoidea. In: Dominick RB, et al. (eds)
The moths of America north of Mexico, including
Greenland. Fasc. 21. E. W. Classey and R. B. D. Publish-
ers, London, 158 pp, 14 pl
Kitching IJ, Cadiou JM (2000) Hawkmoths of the world: an
annotated and illustrated revisionary checklist (Lepi-
doptera: Sphingidae). Cornell University Press, Ithaca,
226 pp, 8 pl
Pinhey ECG (1962) Hawk moths of Central and Southern
Africa. Longmans, Salisbury, 139 pp, 17 pl
Pittaway AR (1993) Hawk moths of the western palaearctic.
E. W. Classey, London, 256 pp
Seitz A (1911–1934) Familie: Sphingidae. In: Die Gross-
Schmetterlinge der Erde, 2:229–273, pl 36–43 (1911–12);
2(suppl): 137–166, 286, pl 12–13 (1932–34); 6:839–900,
pl 90–98 (1931); 10: 23–576, pl 56, 60–68 (1928–29);
14:353–386, pl 61–67 (1927–28). A. Kernen, Stuttgart
Hazelnut and Walnut Twig Borer,
Oberea linearis L. (Coleoptera:
Cerambycidae)
Minos E. Tzanakakis
Aristotle University of Thessaloniki, Thessaloniki,
Greece
The adult (Fig. 8) of this hazelnut and walnut twig
borer is long and slender, measuring 11–16 mm long
and 2 mm wide. The body is black and the legs
yellowish. The antennae do not reach the tip of
the elytra. The pronotum and the elytra are covered
 Hazelnut and Walnut Twig Borer, Oberea linearis L. (Coleoptera: Cerambycidae)
Hazelnut and Walnut Twig Borer, Oberea
­linearis L. (Coleoptera: Cerambycidae),
­Figure 8 Oberea linearis adult.
Hazelnut and Walnut Twig Borer, Oberea
­linearis L. (Coleoptera: Cerambycidae),
­Figure 9 Oberea linearis larva in its gallery in
a ­hazelnut twig.
by fine dark brown hairs. The larva (Fig. 9) is
apodous, ­yellowish or light brown with a brown pro-
thoracic shield, and has the characteristic form of
cerambycid larvae. Its final length is 20–25 mm.
On hazelnut, in southern and central Europe,
Oberea linearis completes one generation every 2
years, whereas in colder regions one in every 3 years.
There is a report that on walnut, in Greece, it com-
pletes one generation per year, but this needs confir-
mation. On hazelnut, this borer spends the first
winter as a relatively young larva and the second
winter as a fully-grown larva at the end of its gallery
in a twig. It pupates in spring. The female lays an egg
in a slit she gnaws in the bark of a shoot, 10–15 cm
from its tip. The first-instar larva bores a gallery
H 1773
which is first semicircular and perpendicular to the
longitudinal axis of the shoot. This causes the ulti-
mate death of the apical part of the shoot which often
breaks off. Subsequently, the gallery goes deeper, and
is directed towards the base of the shoot. The larval
excrement is pushed out through small holes the
larva opens along its gallery. After its first overwin-
tering, the larva continues boring its gallery during
spring and summer of the second year, reaches full
growth in autumn and overwinters. On hazelnut, the
larval gallery reaches 40–60 cm the first year and is
directed towards the base, whereas in the second year
it is directed towards the apex of the twig.
On walnut, in southern Europe, according to
Anagnostopoulos, the adults emerge from the
twigs in May or early June. After feeding on new
growth for a few days and mating, the female lays
an egg on the bark of a shoot a few cm from the tip,
or on the pedicel of a fruit. She prefers shoots bear-
ing fruits, and shoots not very turgid. If the shoot is
turgid, the female gnaws a semicircular to circular
groove in the bark. This weakens the shoot and
seems to favor the development and survival of the
young larva. When the egg is deposited on the fruit
pedicel, the gallery is bored first in the pedicel, then
extended in the shoot. Sometimes the gallery goes
to the mesocarp of the walnut. On walnut, galleries
are shorter than on hazelnut. Galleries in the fruit
pedicels cause early drying and early drop of wal-
nuts, while galleries in the shoots cause the death
of the eroded part of the shoot and of the fruits it
bears. Therefore, on walnut, there may be loss of
fruits, loss of dormant floral buds, and of some
parts of the foliage. The damage may be serious if
the insect’s population density is high. In hazelnuts,
the damage is limited to shoots and twigs and usu-
ally is not such as to justify control measures. Yet,
even on hazelnut economic damage may occur
under excessive population densities of the insect.
On hazelnut, the best method of control is the
removal and destruction of infested dry twigs from
late autumn to early spring, before the adults come
out of them. Another measure is spraying with an
organic contact insecticide of long residual action
when the period the adults come out in spring. On
1774
H Head of Hexapods

walnut, where cutting off infested twigs is usually Head Lice, Pediculus humanus
not practical because of the height of the trees, capitus DeGeer (Phthiraptera:
insecticidal sprays are usually recommended. Sprays Pediculidae)
applied against the codling moth may suffice. Where
pruning is feasible, Anagnostopoulos recommends Head lice are a continuing pest problem for
that it be done in July, to prevent further injury, or humans, especially children.
in spring, when new growth has started and infested  Human Lice
twigs having no leaves can easily be distinguished.  School IPM

References

Balachowsky AS (1962) Entomologie Appliquée a l’ Agricul- Head of Hexapods

ture. Tome I, vol 1. Masson et Cie, Paris, France
Paillot A (1933) Un cerambycide parasite de noyers, Oberea severiano f. gayubo
linearis. Ann Epiphyties 19:369–379
Tzanakakis ME, Katsoyannos BI (1998) Insects of fruit trees Universidad de Salamanca, Salamanca, Spain
and grapevines. Agrotypos, Athens, Greece (in Greek)
In the insects, the head is the tagma in the cephalic
or anterior position. Structures related to two vital
Head functions are situated in this tagma: feeding
(mouthparts) and sensory function (organs like
The anterior body region bearing the eyes, anten-
the antennae and visual organs: two compound
nae (Figs. 10 and 11) and mouthparts.
eyes in the lateral position and three ocelli). The
 Head of Hexapods

Lateral ocellus
Compound eye
Postocular area
Cervix
Gena

Cervical sclerites
Antenna Tentorial suture
Basimandibular sclerite
Maxilla
Clypeus Labium

Labrum Labial palpus

Mandible

Maxillary palpus

Head, Figure 10  Side view of the head of an adult grasshopper, showing some major elements.

Ocelli
Compound eye
Antenna
Head, Figure 11  Anterior aspect of the head and mouthparts of a cicada (Hemiptera).
insertion of the antennae in the head capsule
(Figs. 12 and 14) can vary from the median frontal
area to a more or less ventral position. The com-
pound eyes are situated laterally and generally
occupy an ample surface area. The ocelli are situ-
ated between the compound eyes, arranged like an
isosceles triangle; of the three, the most anterior
(median posi­­tion) seems to be conserved like a
well-differentiated lens, while the two posterior
ones (lateral position) tend to be reduced or modi-
fied, as occurs in some genera of Hymenoptera.
The foramen magnum (Fig.  13) is found in
the posterior part of the head, connecting the head
and the thorax, and on its ventral border is situ-
ated the oral cavity, from which the mouthparts
articulate. It is formed by the acron and the prean-
tennal segment (some consider one labral
­segment), the antennal segment, the intercalary or
premandibular segment (some consider a postan-
tennal fused to the antennal segment), the man-
dibular segment, the maxillary segment, and the
labial segment. Externally, this segmentation is not
visible due to the existence of the head capsule,
formed by the strong sclerotization of the integu-
ment. However, this scelerotization is not homo-
geneous, being differentiated into sutures and
grooves that separate sclerites. Among these
sutures are found the epistomal or frontoclypeal
that unites the anterior tentorial pits and separates
the labroclypeal (preoral) region from the frontal
Head of Hexapods
H 1775
Anterior tentorial pit
clypeus
Mandibular plate
Anteclypeus
Maxillary plate
Labrum
Labium
Mandible
Maxilla
region (postoral); likewise, it separates the frons
from the clypeus. The clypeus is separated from
the labrum by the labroclypeal suture. In certain
species the clypeus is differentiated into an ante-
clypeus and a postclypeus.
The posterior limit of the frons (Fig.  14) is
marked by the branches of the epicranial suture
(in the form of a Y). This suture represents the
ecdysial line of the preimaginal stages, and its
trajectory can vary; thus, postfrontal sutures
appear (delimiting a postfrons) situated exter-
nally at the antennal sockets, and frontal sutures
that are internal to those sockets (delimiting the
frons). Both sutures can coexist, or only one of
them may be present. This generalized arrange-
ment of sclerites and sutures may exhibit numer-
ous modifications, depending on the group of
insects in question.
The odd branch of the epicranial suture,
known as the coronal suture, separates two pari-
etal sclerites that constitute the laterodorsal regions
of the head capsule. The most dorsal zone is called
the vertex (Figs. 14 and 15). Each parietal sclerite
continues ventrally, forming the postocular area
and the gena (areas situated ventrally and posteri-
orly to the compound eye).
Two transverse sutures are generally differen-
tiated in the posterior part of the head capsule: the
occipital and the postoccipital, between which
are found the occiput (dorsal) and the postgenae
1776
H Head of Hexapods

Lateral ocellus
Compound eye

Postocular area
Cervix
Gena

Cervical scleites
Antenna Tentorial suture
Basimandibular sclerite
Maxilla
Clypeus Labium

Labrum Labial palpus

Mandible

Maxillary palpus

Head of Hexapods, Figure 12  Side view of grasshopper head (Orthoptera: Romaleidae).

Occiput

Foram magnum

Posterior ten Occipital condye

tentorial bridge
Maxillary suture
Postgena Posterior tentorial pit

Maxilla

Labium

Maxillary palpus
Labial palpus

Head of Hexapods, Figure 13  Rear view of grasshopper head (Orthoptera: Romaleidae).
 Head of Hexapods
H 1777

Vertex Flagellum
Antenna
Fronts Pedicel
Scape

Mediam ocellus

Face
Anterior tentorior pit
Basimandibular sclerite
Frontoclypeal suture Mandible
Clypeus
Clypeolabral suture
Labrum
Maxillary palpus
Maxilla
Labial palpus

Head of Hexapods, Figure 14  Front view of grasshopper head (Orthoptera: Romaleidae).

Vertex
Antenna
Occipital foramen
Ocellus Compound eye
Occipital condyle

Posterior tentorial bridge

Anterior
tentorial pit
Cardo

Mentum

Clypeus

Labrum Maxillary palpus

Galea Lacinia
Mandible
Labial palpus

Head of Hexapods, Figure 15  Front (left half ) and rear (right half ) view of scorpionfly head (Mecoptera).
1778
H Head of Hexapods
(lateral). The postocciput forms the posterior
margin of the cranium by way of a narrow band
posterior to the postoccipital suture.
The subgenal suture separates the gena and
postgena from a well sclerotized band in the ventral
position, the subgena. The subgenal suture consists
of two parts, an anterior (pleurostomal suture) and
a posterior (hypostomal suture), which delimit the
pleurostoma (in which are found the mandibular
articulations) and the hypostoma (where the max-
illa articulates). The joining of the pleurostoma and
hypostoma forms the subgena. Certain authors uti-
lize the term epicranium to denote what together is
formed by the occiput (posterior), vertex (dorsal),
frontoclypeus (anterior) and genae (lateral).
Internally, the head has some endoskeletal
structures, the occipital phragma and the tento-
rium. The occipital phragma corresponds to an
internal ridge or phragma of the postoccipital
suture. This phragma normally is formed by a pair
of laterodorsal plates, in which the gnathal mus-
cles and the prothoracic motor muscles of the
head are inserted. The tentorium is a complex
structure that has two fundamental functions;
it serves as the area of muscular insertion and
Dorsal tentorial arm
Tentorial ridge
Anterior tentorial arm
Subgenal ridge
Epistomal ridge
Anterior tentorial pit
Epistomal suture Clypeus
Labrum
Head of Hexapods, Figure 16  Internal structures of the insect cranium (after Snodgrass).
reinforces the head capsule. The number and type
of muscles that are inserted into the tentorium
varies depending on the hexapod group. In primi-
tive groups of hexapods like the Machilidae it con-
sists of three parts, the anterior and posterior
tentoria and the tentorial-neck plate.
In the Pterygota, the tentorium is an odd
structure (Fig. 16) formed by the tentorial body,
from which are differentiated three pairs of exten-
sions, one pair in the anterior position (anterior
arms or pretentorium), another posterior (poste-
rior arms or metatentorium) and a third dorsal
(dorsal arms or supratentorium). Anterior and
posterior arms result from anterior invaginations
(pretentorinae) and posterior invaginations
(metatentorinae), respectively. They are apodemal
structures. These invaginations constitute impor-
tant morphological reference points. The pair of
dorsal arms is directed toward the antennal region.
Their union with the cuticle is through naturally
fibrous connective structures. Those dorsal arms
constitute evaginations of the anterior arms.
Two types of head capsules are distinguished,
depending on whether the mouthparts are posi-
tioned externally (ectotrophous or ectognathous) or
Occiput
Postoccipital ridge
Postoccipital suture
Posterior tentorial pit
Subgenal suture
Anterior tentorial pit
Subgenal suture

internally (entotrophous or entognathous). These
terms can also be applied to the mouthparts them-
selves. In the first case, both the mouthparts and
their articulations with the head are visible exter-
nally. In the case of the entognathous insects, the
mouthparts are withdrawn into the head, in internal
cavities, and are not visible externally when the
insect is at rest. Clear examples of this type of head
capsule are found in Collembola, Protura and Diplura,
and to a lesser extent in ­Thysanoptera (Fig. 20), Phthi-
raptera (Anoplura) and some Hemiptera (Fig. 17).
If we consider the position of the oral cavity
and consequently of the mouthparts with respect
to the cephalocaudal axis, three fundamental types
of head capsules (Fig. 21) are distinguished: orthog-
nathous or hypognathous; prognathous; and opist-
hognathous or opisthorhynchous (these names can
also be applied to the mouthparts).
In the orthognathous or hypognathous condi-
tion, the mouthparts are situated more or less per-
pendicular to the cephalocaudal axis. This model is
differentiated in phytophagous insects that feed
on  solid particles and in some predators. In the
prognathous condition the oral cavity is situated
in  the anterior ventral part of the head, and the
mouthparts are directed forward. Modifications of
certain parts of the head capsule result, such as a
tendency to incorporate the clypeus into the frons,
Ocelli
Compound eye
Antenna
Head of Hexapods, Figure 17  Front view of cicada head (Hemiptera: Cicadidae).
Head of Hexapods
H 1779
a shortening of the occipital areas, and a lengthen-
ing of the structures that form the ventral closure of
the head. The prognathous condition is represented
in many hexapod orders, being considered a primi-
tive condition (for example, Embiidina, Isoptera
and Dermaptera) or secondarily acquired (larvae of
Neuroptera, mining caterpillars and certain beetle
larvae). The prognathous condition is characterized
by the predatory species, whose adults actively
hunt their prey. The condition is also found in some
larval forms (particularly in  Coleoptera) that use
their mandibles for burrowing.
The opisthognathous condition is character-
ized by the oral cavity being (Fig. 17) situated in the
posterior ventral part of the head and the mouth-
parts directed toward the back, being held at rest
between the anterior legs (Hemiptera).
There exists a tendency for the ventral closure
of the head to be differentiated into two types of
head or cranial capsules: open and closed. In the
first case, the foramen magnum is bordered ven-
trally by the proximal margin of the submentum,
having the metatentorinae in the inferior extremes
of the occipital suture, near the foramen magnum.
Nevertheless, the most frequent case is where the
labium is separated from the foramen magnum by
a more or less wide, sclerotized band. This band
can be situated anteriorly to the metatentorinae,
Anterior tentorial pit
clypeus
Mandibular plate
Anteclypeus
Maxillary plate
Labrum
Labium
Mandible
Maxilla
1780
H Head of Hexapods

Antenna

Ocellus Compound eye

Clypeus

Epipharynx

Labrum
Labial palpus Galea

Head of Hexapods, Figure 18  Dorsolateral view of a moth head (Lepidoptera).

Compound eye

Antenna

Anterior tentorial pit

Clypeus
Base of mandible
Mentum Unsclerotized labrum
Sclerotized labrum
Maxillary palpus
Labellum of labium Maxilla

Head of Hexapods, Figure 19  Front view of a horse fly head (Diptera: Tabanidae).

forming the hypostomal or postgenal bridge, or it
can be posterior to the metatentorinae, forming
the gula. These sclerotized formations can occupy
a great part of the ventral zone of the head, with
the consequent displacement of the mouthparts
(Fig. 21) toward the front. The prognathous condi-
tion discussed earlier originated in this manner.
The hypostomal bridge is formed by the
union of two lobes that are of mixed origin (each
one is formed beginning from the hypostoma and
the postgena). Clear examples of this bridge exist
in advanced Hymenoptera and Diptera.
The gula constitutes a sclerotized zone that
prolongs the submentum behind the metatento-
rinae, which are advanced, and it is separated
from the postgenae by the gular sutures. The ori-
gin of the gula is disputable, and although it
probably pertains to what is called labial, some
attribute its origin starting from the neck and
including the prothoracic region. Clear examples
of the presence of the gula are found in certain
beetles.
 Antennae of Hexapods
 Mouthparts of Hexapods
 Head of Hexapods
H 1781

Antenna

Compound eye

Anterior tentorial pit

Transverse clypeal suture
Clypeus

Labrum
Maxilla Labium
Labial palpus
Maxillary palpus

Head of Hexapods, Figure 20  Front view of a thrips head (Thysanoptera).

Antenna
Antenna Movable hook of palpus
Eye Labrum

Palpus
Eye

Proboscis
Mentum Leg
Maxillary palpus

Head of Hexapods, Figure 21  Comparison of mouthpart orientation: hypognathous (left), prognathous
(center), and opisthognathous (right).

References Imms AD (1957) A general textbook of entomology, 9th edn.

Bitsch J (1973) Morphologie de la tête des insects. Partie
Générale. In: Grassé PP (Dir.) Traité de Zoologie, VIII(I)
19:3–100
Bitsch J (1994) The morphological groundplan of Hexapoda:
critical review of recent concepts. Ann Soc Entomol
France 30:103–129
Borror DJ, Triplehorn CA, Johnson NF (1989) An introduc-
tion to the study of insects. Saunders College Publish-
ing, Philadelphia, Pennsylvania
Gillot C (1995) Entomology, 2nd edn. Plenum Press,
New York, NY
Methuen, London, UK
Manton SM (1977) The Arthropoda. Habits, functional mor-
phology and evolution. Clarendon Press, Oxford, UK
Matsuda R (1965) Morphology and evolution of the insect
head. Mem Am Entomol Inst 4:1–334
Nauman ID (ed) (1994) Systematic and applied entomology.
An introduction. Melbourne University Press, M
­ elbourne,
Australia
Quéinnec E (2001) Insights into arthropod head evolution.
Two heads in one: the end of the “endless dispute?” Ann
Soc Entomol France 37:51–70
Richards OW, Davies RG (1977) Imms’ general textbook of
entomology, 10th edn. Chapman and Hall, London, UK
1782
H Heart

Snodgrass RE (1951) Comparative studies on the head of  American Butterfly Moths

mandibulate arthropods. Comstock Publishing, Ithaca,
 Butterflies and Moths
New York
Snodgrass RE (1952) A textbook of arthropod anatomy.
Comstock Publishing, Ithaca, New York
Steinmann H, Zombori LM (1981) An atlas of insect mor- Heel Flies
phology. Akademiai Kiado, Budapest, Hungary

Members of the family Oestridae (order Diptera).

 Flies (Diptera)
Heart  Myiasis

Also known as the dorsal vessel (Figs. 22 and 23),

it is a muscular tube extending dorsally through
the abdomen of  the insect which pushes blood
(hemolymph) forward into the aorta and to
the head.
Hebridae
A family of bugs (order Hemiptera). They some-
times are called velvet water bugs.
 Bugs
Hedylidae
A family of moths (order Lepidoptera) also known
as American butterfly moths.
Heer, Oswald
Oswald Heer was born in Matt, a mountain village
in Switzerland, on August 31, 1809. His interest in
natural history was aroused there. In 1828, he went
to Universität Halle in Germany to study theology
(his father was a priest) and there met several
renowned naturalists. He was ordained in St. Gallen,
Switzerland, but also pursued natural history. In
1832, he obtained the post of curator of the Escher-
Zollikofer natural history collection in Zurich. Two
years later he became docent of botany and zool-
ogy of Universität Zurich, and in 1835 professor of
botany there. His work on paleobotany and paleo-
zoology was perhaps his most notable research
contribution. Nevertheless, his multi-part “Fauna
Coleopterorum Helvetica” was seminal for Swiss

Dorsal Pulsatile
Ostia Heart diaphragm organ Aorta

Antennal
pulsatile organ

Cerebral ganglion

Ventral Nerve Septa

diaphragm cord

Heart, Figure 22  Diagram of the components of the insect circulatory system; hemolymph moves
through the heart and aorta to the head, filtering back through the body cavity and back into the heart
via the ostia. Pulsatile organs assist in moving hemolymph through the appendages, especially the
wings. (adapted from Chapman, The insects: structure and function).
 Helicosporidium
H 1783

Heart Helicosporidium
Muscle aurélien tartar
University of Florida, Gainesville, FL, USA
Silk gland

Midgut To date, there is only one named species of Heli-

cosporidia: Helicosporidium parasiticum. It was ini-
Visceral fat tially described and named by Keilin in 1921, who
Parietal fat
detected this protist in larvae of Dasyhelea obscura
Winnertz (Diptera: Ceratopogonidae) collected in
Nerve cord England. He examined the new parasite thoroughly
and attempted to infer its life history from his
observations. He described a vegetative growth
Heart, Figure 23  Cross section of caterpillar larva characterized by very active multiplications of heli-
showing the location of the heart (adapted from cosporidial cells inside the body cavity of the host,
Chapman, The insects: structure and function). and noticed that these “schizogonic multiplica-
tions” were followed by the formation of what he
entomology, and is still consulted. He died on called spores. Keilin noted that the spores are very
­September 27, 1883, in Lausanne. easily recognized; they consist of the assembly,
inside an external membrane, of three ovoid cells
(named by Keilin “sporozoites”) and one periph-
Reference eral, spiral, filamentous cell. These features, espe-
cially the highly characteristic filamentous cell,
Herman LH (2001) Heer, Oswald. Bull Am Mus Nat Hist have since remained the principal diagnostic for
265:76–77 identification of a Helicosporidium sp. Keilin was
able to describe and characterize structurally the
new genus Helicosporidium, and the new species
Heleomyzidae H.  parasiticum. He also was able to present a
hypothetical life cycle of this protist based on his
A family of flies (order Diptera). They commonly microscopy observations. He suggested that the
are known as heleomyzid flies. spores (or cysts) break open in the host hemocoel,
 Flies releasing the filamentous cell and the three “sporo-
zoites,” which he proposed are the infective
forms of H. parasiticum (Fig. 24). Keilin believed
Helgrammite that H. parasiticum belonged to the Protozoa, and
compared this isolate with members of various
The aquatic larvae of corydalids (Megaloptera: clades: Cnidiosporidia (which, at that time, included
Corydalidae). Microsporidia such as Nosema bombicis), Hap-
 Alderflies and Dobsonflies losporidia, Serumsporidia, and Mycetozoa. He
concluded that the genus Helicosporidium differed
markedly not only from all these groups, but also
Helicopsychidae from all the protists known at that time. He finally
proposed that Helicosporidium “forms a new group,
A family of caddisflies (order Trichoptera). which may be temporarily be included in the
 Caddisflies group of the Sporozoa.”
1784
H Helicosporidium
Helicosporidium, Figure 24  Light micrograph of dehisced Helicosporidium sp. cysts. Following rupture
of the pellicle, one filamentous cell and three ovoid cells are released from each cyst.
Following the discovery of another isolate
of Helicosporidium parasiticum in a larva of
Hepialis pallens (Lepidoptera: Hepialidae),
another taxonomic position was proposed for
the group Helicosporidia. Based on observation
of this new isolate as well as the original speci-
men described by Keilin, Weiser claimed that the
Helicosporidia are best placed among the lower
Fungi. He argued that the spore characteristics
are much too different from what is found in
Protozoa, but are similar in some aspects to
primitive Fungi, such as insect pathogens of the
genus Monosporella, classified as Nematosporoi-
deae inside the Saccharomycetaceae (primitive
Ascomycetes).
In the early 1970s, Helicosporidium parasiti-
cum was isolated from larvae and adults of the
beetle Carpophilus mutilatus (Coleoptera: Niti-
dulidae). This isolate was infectious per os to 18
species of arthropods, including three orders of
insects (Lepidoptera, Coleoptera, Diptera) and
one family of mites. In contrast, species of
Orthoptera, Hymenoptera, and Diptera were not
susceptible to this isolate. The spores (cysts)
present in the host artificial diet were ingested
and released the three round cells and the fila-
mentous cells in the host midgut. After 24 h,
helicosporidial cells appeared in the host hemo-
lymph and grew vegetatively. The vegetative
growth is characterized by cell division that
occurs within a pellicle. After division, the pelli-
cle ruptures and releases the daughter cells (4 or
8). Empty pellicles and daughter cells eventually
fill the entire host hemocoel. Daughter cells then
develop into spores, in which the filamentous
cell differentiates and encircles the three round
cells. Additional studies reported the presence
of  Helicosporidium sp. in new host species
such  as  crustaceans, mites, collembolans, and
trematodes.
In 1999, a Helicosporidium sp. was discov-
ered in larvae of the black fly Simulium jonesi
Stone & Snoddy (Diptera: Simuliidae). This iso-
late was demonstrated to grow under both in
vitro and in vivo conditions. In vivo, the ingested
helicosporidial cysts dehisced and released the
three ovoid cells and the filamentous cells. The
filamentous cells attached to the peritrophic
membrane and penetrated the midgut epithelium
initiating infection. The in vitro growth of Heli-
cosporidium sp. was reminiscent of that reported
for unicellular, achlorophytic algae belonging to
the genus Prototheca. Both the genera Helicospo-
ridium and Prototheca are characterized by a veg-
etative growth that consists of cell divisions
inside a membrane. Four, eight, or sixteen daugh-
ter cells are produced inside this pellicle and
eventually are released. Such cell divisions result
 Hemelytron (pl. hemelytra)
H 1785

in the accumulation of both round daughter cells Heliozelidae

and empty pellicles. The suggestion that Heli-
cosporidium was related to algae has been con- A family of moths (order Lepidoptera). They com-
firmed by molecular analysis. The 18S, 26S, 5.8S monly are known as shield bearer moths.
regions of the Helicosporidium ribosomal DNA,  Shield Bearer Moths
as well as some partial sequences of the actin  Butterflies and Moths
and tubulin genes, were sequenced. Comparative
analyses of these nucleotide sequences were
­performed in order to evaluate the position of Heloridae
Helicosporidium sp. within the phylogeny of
eukaryotes. All trees depicted Helicosporidium A family of wasps (order Hymenoptera).
sp.  as a green alga (Chlorophyta), and as sister  Wasps, Ants, Bees and Sawflies
taxon to the genus Prototheca (class Trebouxio-
phyceae). This association always was supported Helosciomyzidae
by significant bootstrap values. On the basis of this
phylogenetic analysis, Helicosporidium sp. is clearly A family of flies (order Diptera).
neither a protozoan nor a fungus, but represents  Flies
the first described algal invertebrate pathogen.

Helotrephidae
References
A family of bugs (order Hemiptera).
Boucias DG, Becnel JJ, White SE, Bott M (2001) In vivo and  Bugs
in vitro development of the protist Helicosporidium sp.
J Eukaryotic Microbiol 48:460–470
Kellen WR, Lindegren JE (1974) Life cycle of Helicosporidium
parasiticum in the navelworm Paramyelois transitella. Hematophagous
J Invertebr Pathol 23:202–208
Tartar A, Boucias DG, Adams BJ, Becnel JJ (2002) Phyloge-
netic analysis identifies the invertebrate pathogen Heli-
(haematophagous) Insects that feed on blood.
cosporidium sp. as a green alga (Chlorophyta). Int J Syst
Evol Microbiol 52:273–279
Hematophagy
Heliodinidae (haematophagy) Feeding on blood. Usually this
refers to arthropods that bite to obtain blood, but
A family of moths (order Lepidoptera). They com- secondary hematophagy also is possible, wherein
monly are known as sun moths. arthropods feed at a bleeding wound, with the
 Sun Moths wound accidental or inflicted by another organism.
 Butterflies and Moths

Hemelytron (pl. hemelytra)

Heliomyzid Flies
The front wing of insects in which  the basal
Members of the family Heliomyzidae (order portion in thickened and the ­distal portion mem-
Diptera). branous (Fig. 25), usually Hemiptera.
 Flies  Wings of Insects
1786
H Hemerobiidae

Embolium Fracture

R Cuneus
Sc
r-m
M M M
R-M Membrane
Cu Corium

P Cu
1A 1A
Clavus

Anal ridge

Hemelytron (pl. hemelytra), Figure 25  Front wing of a bug (Hemiptera: suborder Heteroptera), thickened
basally and membranous distally.

Hemerobiidae modified form of hemimetabolous development are

A family of insects in the order Neuroptera. They
commonly are known as brown lacewings.
 Lacewings, Antlions and Mantidflies
Hemipsocidae
A family of psocids (order Psocoptera).
 Bark-Lice, Book-Lice or Psocids
said to have paurometabolous development. The
hemimetabolous insects (including paurometabo-
lous insects) lack the pupal stage that is found among
insects with holometabolous development. The
immature forms of hemimetabolous insects are
called nymphs, though some entomologists call
all  immature insects larvae, regardless of develop-
mental pattern. (contrast with holometabolous
development)
 Metamorphosis
 Complete Metamorphosis
 Incomplete Metamorphosis

Hemiptera
An order of insects. Many of these insects are
called bugs, or true bugs, but this is a very diverse
assemblage, and includes leafhoppers, treehop-
pers, cicada, scales, aphids, whiteflies, and many
others. Hemiptera is sometimes treated as two
orders: Hemiptera and Homoptera.
 Bugs
Hemimetabolous Development
Insects having incomplete metamorphosis, or a
gradual change in body form with each molt, and
wings developing externally. A subset of insects
(Odonata, Ephermeroptera, Plecoptera) with a
Hemocoel
(haemocoel) The hemolymph (blood)- filled body
cavity of arthropods.
Hemocoelic Insemination
Female bed bugs (Hemiptera: Cimicidae) are
inseminated by the male’s needle-like penis, which
can be inserted in many locations in the female’s
body, because the sperm migrate through the
hemolymph to the ovary. This process may provide
additional nutrients to the female, but certainly
other insects do this in less traumatic ways. This
process is also called “traumatic insemination.”
 Bed Bugs
 Hemocytes of Insects: Their Morphology and Function
H 1787

Hemocytes of Insects: Their encapsulation process in Lepidoptera, granulo-

Morphology and Function cytes were determined to attach to, and release
their contents (degranulate) on, the surface of a
pauline o. lawrence foreign object during the initial phase of capsule
University of Florida, Gainesville, FL, USA formation, and to release cytokines that recruit
plasmatocytes (Fig. 26) to the site. The innermost
Hemocytes are blood cells that circulate in a clear cells of the capsule then flatten and develop
fluid, the plasma, within the hemocele (body cav- ­desmosomes that hold the cells together. Gap-
ity) of insects. The hemocytes, plasma, and dis- junctions develop between cells of the external
solved inorganic and organic molecules constitute and internal layers of the capsule, providing an
the hemolymph. Nutrients, nitrogenous waste intercellular pathway presumably for nutrients,
products, hormones, and a variety of other sub- hormones, and probably cytokines for intercellu-
stances also are transported in the hemolymph lar signaling, and for other factors needed to main-
for distribution to various tissues. In most insect tain the integrity of the capsule. Granulocytes are
larvae, the hemocytes are produced within presumed to arise from plasmatocytes, and their
hematopoietic (hemopoietic) organs (blood cell
producing factories) where they replicate and dif-
ferentiate. The number and types of hemocytes
vary with insect species, developmental stage, and
physiological state. In general, total hemocyte
count (the number of hemocytes per unit volume)
increases throughout larval development, reach-
ing a maximum at each larval ecydsis but probably
not during pupal/adult ecdysis. Qualitative and
quantitative fluctuations in hemocyte populations
also may be influenced by endocrine events as well
as by wounding and microbial and parasitic
infections.
The classification of insect hemocytes has
been based primarily on the morphology of cells
observed under the light and electron micro-
scopes. While all hemocyte types described to date
do not occur in all insects, the six recognized and
most common types are: adipohemocytes, granu-
locytes, oenocytoids, plasmatocytes, prohemo- Hemocytes of Insects: Their Morphology and
cytes, and spherule cells. Adipohemocytes vary in ­Function, Figure 26  Transmission electron
size and are ovoid with well-defined lipid droplets ­micrograph of a spherical (discoid) plasmatocyte
and granular cytoplasm with rough endoplasmic (a) from the third instar of the Caribbean fruit
reticula (RER) and Golgi complexes, indicative of fly, Anastrepha suspensa (Diptera: Tephritidae).
synthetic and secretory functions. Granulocytes Three populations of discoid plasmatocytes
vary in size and shape and have a small nucleus: occur in A. suspensa, ranging from 10–25 μm in
cytoplasm ratio and small, uniform acidophilic ­diameter. (b) Portion of a lamellocyte showing
cytoplasmic inclusions. Their cytoplasm is richly rough e
­ ndoplasmic r­ eticula (RER). Lamellocytes
supplied with an elaborate network of RER, Golgi are 15–20 μm long and 6–10 μm wide. N = nucleus.
vesicles and other organelles. In studies of the (Micrograph by P.O. Lawrence.)
1788
H Hemocytes of Insects: Their Morphology and Function
population ranges from 30 to 65% of the total
hemocyte count. Some authors consider cysto-
cytes (also called coagulocytes) to be a small form
of granulocytes because they are presumed to par-
ticipate in the coagulation process. Like granulo-
cytes, cystocytes have acidic cytoplasmic inclusions
and a small nucleus. Because their cell cytoplasm
is translucent, cystocytes sometimes are called
hyaline cells. To date, cystocytes have not been
identified in lepidopterans, dipterans or
hymenopterans.
Oenocytoids are large (18–38 μm in diame-
ter), sometimes with two eccentric nuclei and
an  acidophilic cytoplasm. They have numerous
microtubules and few synthetic organelles. They
also exhibit endogenous phenoloxidase activity
and are presumed to participate in the process of
melanization (melanosis), often associated with
wound healing and encapsulation. Plasmatocytes
usually represent more than 28% of the total
hemocyte population in most insects. They are
pleiomorphic, varying in size and shape from
round (discoid), to spindle-shaped (lamellocytes)
and have a single nucleus that occupies about
40–50% of the granular, basophilic cytoplasm.
They possess RER and Golgi organelles, indica-
tive of a synthetic function. They are motile and
phagocytic, with lysosomal vesicles, and are the
primary line of defense in the insects’ cellular
response to microbial infection. Indeed, they
release substances that induce degranulation of
the granular hemocytes during the final stage of
encapsulation. They also participate in other
defense-related ­processes such as nodulation
and, along with ­cystocytes, in plasma coagula-
tion. Podocytesare considered by some authors
to either arise from, or be a form of, plasmato-
cyte. Prohemocytes are thought to be the small-
est blood cells. They are round with a large
nucleus:cytoplasm ratio, and basophilic, but are
devoid of synthetic organelles (e.g., endoplasmic
reticula, etc). They undergo mitosis, presumably
giving rise to prohemocytes, plasmatocytes, and
probably to granulocytes. Thus, they are consid-
ered the hemocyte stem cells. Spherulocytes
(spherule cells) are ovoid and usually are larger
than prohemocytes, but smaller than some plas-
matocytes. In some cases, spherulocytes repre-
sent about 4% of the total hemocyte population.
Their cytoplasm appears to contain large
­acidophilic inclusions. Other hemocytes such as
podocytes and vermiform cells could be forms of
plasmatocytes, but neither their origins nor spe-
cific functions are known.
Larval dipterans have hemocytes (e.g., pro-
hemocytes, plasmatocytes, and oenocytoids)
similar to some found in other insects. However,
they are thought to lack true granulocytes and
spherule cells. Indeed, some authors consider
previously described “spherule cells” and “granu-
locytes” of some Diptera (e.g., Calliphora erythro-
cephala) to be plasmatocytes and thrombocytoids,
respectively. True spherule cells are thought to be
absent in ­dipterans while thrombocytoids, crys-
tal cells, and vacuolated cells are presumed to be
unique to ­dipterans, although they do not occur
in all species.
Crystal cells, unique to drosophilids, are
10–12.2 μm with a nucleus of 3.5–5 μm in diameter
and comprise 5–10% of the total hemocyte count.
They characteristically have large cytoplasmic para­
crystalline inclusions that are presumed to contain
tyrosine, a substrate of phenol oxidase. Conse-
quently, upon disruption, these cells release their
contents, causing melanosis which is important in
wound healing and other defense responses. Since
oenocytoids of other dipterans, and indeed other
insects, are known to have endogenous ­phenol
­oxidase activity and are involved in wound healing,
it is likely that the crystal cells of Drosophila are at
least analogous, if not homologous to them.
Thrombocytoids are round cells with numer-
ous invaginations of the plasma membrane that
penetrate deep into the cytoplasm close to the
nucleus. Based on in vitro studies, thrombocytoids
are presumed to phagocytose bacteria, and to
­participate in encapsulation and wound healing.
They disintegrate within the hemolymph to give
rise to cytoplasmic fragments and naked nuclei.
Thrombocytoids are absent from Drosophila sp.

and culicid mosquitoes, but occur in calliphorids
and tipulids.
Vacuolated cells (Fig.  27) are 14–23 μm in
diameter and are characterized by extensive
­vacuolation of the cytoplasm, presumably the
result of invaginations of the cell membrane.
Numerous free ribosomes, RER, Golgi bodies,
and mitochondria are dispersed throughout the
cytoplasm. Vacuolated cells are <20% of the total
hemocyte count. These cells are presumed to be
unique to the Diptera, although do not occur in
all species. They have been reported from larval
sciarids, the fruit fly ­Anastrepha ­suspensa, and
from mutant Drosophila melanogaster that have
a lethal malignant blood neoplasm (lmbn) (i.e.,
­melanotic tumors of the hemocytes). These
tumerous blood cells [l(2)mbn] when cultured
in vitro, readily differentiate into hemocyte mor-
photypes that correspond to plasmatocytes,
Hemocytes of Insects: Their Morphology and
­Function, Figure 27  Transmission electron
­micrograph of a portion of a vacuolated ­hemocyte
from the third instar of the Caribbean fruit fly,
Anastrepha suspensa. These vacuolated cells are
25–30 μm in diameter (compared to 14–23 μm
in other dipterans). N = nucleus, V = vacuoles.
­(Micrograph by P.O. Lawrence.)
Hemocytes of Insects: Their Morphology and Function
H 1789
lamellocytes, podocytes, and vacuolated cells of
the wild type (normal) larvae. Lamellocytoids
(apparent homologs of the lamellocytes in vivo)
were transformed to vacuolated-like cells in
vitro upon treatment with the insect molting
­hormone 20-hydroxyecdysone (20-OHE). Inter-
estingly, the combined application of juvenile
hormone III (the predominant juvenile hormone
found in dipterans) and related JH agonists,
along with 20-OHE, counteracted the effects of
20-OHE alone. Given the respective roles of JH
and 20-OHE in the maintenance of larval char-
acteristics and induction of metamorphosis, it is
likely that the incidence of vacuolated cells in
dipterans may be influenced by hormonal and/
or related developmental events, as is apparently
the case with certain other hemocytes.
The behavior and function of hemocytes are
influenced by the release of humoral factors such
as cytokines, cell adhesion molecules, lectins, and
agglutinins that mediate intercellular communica-
tion. The receptors for these factors have not been
fully characterized, and neither is their distribu-
tion among the different hemocytes.
Overall, it appears that certain hemocytes,
such as the plasmatocytes, have similar morphol-
ogy and ultrastructure across the various insect
families. Indeed, techniques have been used to
establish criteria that facilitate interspecies com-
parisons and classification of some hemocytes.
These include the use of electron microscopy, the
development of monoclonal antibodies for spe-
cific hemocytes, and histochemistry. Nevertheless,
additional approaches are needed to identify
unique characteristics of hemocyte types and
facilitate meaningful interspecies and within spe-
cies comparisons. The plasticity in hemocyte
morphology as a result of normal ontogeny, stage
specific factors, and hormonal or environmental
events de­­mands additional approaches to provide
insights into hemocyte identity within and
between species and families. Likely technologies
include differential display, microarrays and
glycobiology.
 Hemolymph
1790
H Hemolymph
References
Brehelin M, Zachary D (1986) Insect haemocytes: a new clas-
sification to rule out the controversy. In: Brehelin M (ed)
Immunity in invertebrates. Springer-Verlag, Berlin,
Germany, pp 36–48
Caveney S, Berdan R (1982) Selectivity in junctional coupling
between cells of insect tissues. In: King RC, Akai H (eds)
Insect ultrastructure. Plenum Press, New York, NY,
pp 434–465
Ratcliffe N (1993) Cellular defense responses of insects: unre-
solved problems. In: Beckage NE, Thompson SN, Federici
BA (eds) Parasites and pathogens of insects, vol 1:
­Parasites. Academic Press, San Diego, CA, pp 267–304
Shrestha R, Gateff E (1986) Ultrastructure and cytochemistry
of the tumorous blood cells in the mutant lethal(3)
malignant blood neoplasm of Drosophila melanogaster.
J Invertebr Pathol 48:1–12
Strand M, Pech L (1995) Immunological basis for compatibil-
ity in parasitoid-host relationships. Ann Rev Entomol
40:31–56
Hemolymph
(haemolymph) The fluid filling the hemocoel
and dorsal vessel (Fig.  28). The blood cells, or
hemocytes, are considered to be part of the hemo-
lymph. The insect’s “blood.” The hemolymph has
several important functions, including the transport
of nutrients to cells and tissues; the transport of
Dorsal
Ostia Heart diaphragm
Ventral Nerve Septa
diaphragm cord
Hemolymph, Figure 28  Diagram of the components of the insect circulatory system; hemolymph moves
through the heart and aorta to the head, filtering back through the body cavity and back into the heart
via the ostia. Pulsatile organs assist in moving hemolymph through the appendages, especially the
wings. (adapted from Chapman, The insects: structure and function).
­ ormones, waste products and water; the transport
h
of body heat away from organs generating the heat,
or the localization of heat in tissues such as tho-
racic muscles where heat is needed; it serves as a
reservoir of nutrients, enzymes and fluids; it func-
tions as a sink for carbon dioxide; it serves as a
lubricant and hydraulic fluid for the maintenance
of body shape and the expansion of wings; the
hemocytes attack invading organisms and foreign
substances.
Hemolytic Anemia
(haemolytic anemia) A shortage of red blood cells
due to their destruction. This sometimes occurs in
the case of envenomization by brown recluse
spider.
Hemophagous
Blood feeding. This term is used to describe biting
flies, fleas, mites, ticks and mites. Many blood-
feeding arthropods vector diseases.
 Vector Capability of Blood Sucking
Arthropods: A Forecasting Matrix
Pulsatile
organ Aorta
Antennal
pulsatile organ
Cerebral ganglion

Henicocoridae
A family of bugs (order Hemiptera, suborder
Pentamorpha).
 Bugs
Henneguy, Louis Félix
Félix Henneguy was born in Paris on March 18,
1850. He studied medicine, was an assistant in
­physiology in Montpellier (1871–1875) and earned
a doctorate in medicine there. He moved to Paris in
1881 and became a preparator in comparative
embryology at the Collège de France. He earned a
doctorate in science in 1888. In 1900 he became a
professor of comparative embryology. In 1907 he
was elected member of the Academie de Médecine,
and in 1908 of the Academie des Sciences. His
other interest, in entomology, led to his (1904) book
“Les insectes” which emphasizes morphology and
­embryology. He also contributed some shorter
­entomological papers. He died on January 28, 1928.
Reference
Howard LO (1928) Obituary. Entomol News 39:136
Hennig, Willi
Willi Hennig was born in Durhennersdorf,
­Germany, on April 20, 1913. As a youth, he collected
beetles and butterflies, was interested in birds and
reptiles, started a herbarium, and liked to visit the
zoological museum in Dresden. In 1932, he entered
Universität Leipzig to study zoology, botany and
geology. He graduated in 1936, having written a
­thesis on the copulatory organ of cyclorraphous
Diptera, and also having published eight papers.
He  trained briefly at the zoological museum in
­Dresden, then went to the Deutsche Entomologische
Institut in January 1937. By 1939 he had published
some 41 papers, most of them on flies. But, in the
Hennig, Willi
H 1791
spring of 1939, he was drafted into the German
army, in which he served as infantryman and was
wounded. Next, he was sent by the German army to
northern Italy, to work in malaria control. Captured
by British troops, he continued to work in malaria
control. Before he was sent home to Germany, he
drafted a manuscript under the title “Grundzüge
einer Theorie der phylogenetischen Systematik.”
Even during the war he published, including about
his work in malaria control. After returning to
­Germany, he spent some months in several cities,
including 13 months at Universität Leipzig, until in
April 1947 he was able to return to the Deutsche
Entomologische Institut in Berlin. He continued to
work on his manuscript on larval forms of Diptera,
which eventually was published (beginning in 1948,
in three volumes) as “Die Larvenformen der Dip-
teren.” He worked some more on his “Grundzüge
einer Theorie…” and then published it in 1950.
Numerous other publications followed, but then the
government of East Germany in 1961 built a wall
across the city of Berlin. Hennig could no longer
travel from his home in West Berlin to the institute
in East Berlin. Instead of emigrating to what was
then the communist country of East Germany, he
resigned from his job, and taught for 2 years at the
Technische Universität in Berlin. Then, he moved
with his family to Ludwigsburg (West Germany) in
1963. He resumed an earlier interest in fossil insects,
and published (1964–1972) 17 papers on Diptera
from Baltic amber and three on fossils from Leba-
nese amber. The insight this gave him resulted in his
(1969) book “Die Stammesgeschichte der Insekten,”
translated (1981) to English as “Insect phylogeny.”
Meanwhile, his “Grundzuge…” had been translated
to English as (1966) “Phylogenetic systematics,”
whose translation and concomitant editing did not
please Willi. A ­German edition, truer to its author’ s
intent, was published (1982) as “Phylogenetische
Systematik.” He continued working on taxonomic
revisions of flies of the palearctic region, and pub-
lished works on the families Muscidae (1955–1964)
and Anthomyiidae (1966–1976) in Lindner’s multi-
volume work “Die Fliegen der palaearktischen
Region,” which were his 13th and 14th contributions
1792
H Hepialidae

to this work. He was appointed associate professor at Herbicide

Universität Tübingen in 1970, and gave lectures
there, and he was co-editor of the journal Zeitschrift A pesticide used to control weeds.
für Morphologie der Tiere/Zoomorphologie. In his
last year, he was working on a revised edition of “Die
Larvenformen der Dipteren.” He had a very impor- Herbivore
tant influence on biology as a whole because of
his phylogenetic theory, even taking into account Animal that consumes plant tissue. A phytophage.
his dissatisfaction at its translation into English. He (contrast with carnivore)
died on November 4, 1976, survived by his always-
supportive wife and their three sons.
Herbivory

References Feeding by insects on plant parts (= phytophagy,

though herbivory can be more narrowly defined
Kuhne WG (1978) Willi Hennig 1913–1976: Die Schaffung as feeding on herbage [grass and other low-growing
einer Wissenschaftstheorie. Entomol Germanica plants, not trees]). Such animals are said to be
4:374–376
Schlee D (1978) In Memoriam Willi Hennig 1913–1976. Eine
herbivorous or herbivores.
biographische Skizze. Entomol Germanica 4:377–391  Phytophagy
 Food Habits of Insects
 Allelochemicals
Hepialidae  Plant Secondary Compounds and
Phytophagous Insects
A family of moths (order Lepidoptera). They com-  Graminivory
monly are known as ghost moths and swifts.  Granivory
 Ghost Moths  Folivory
 Butterflies and Moths

Herman, Ottó
Heptageniidae
george hangay
A family of mayflies (order Ephemeroptera). Narrabeen, NSW, Australia
 Mayflies
Ottó Herman was born on June 26, 1835 at
Reznóbánya, in Zólyom Shire, Hungary. He was
Heptapsogastridae introduced to natural sciences, especially the study
of birds, at an early age, as his father was a medical
A family of chewing lice (order Phthiraptera). practitioner and keen amateur ornithologist. How-
 Chewing and Sucking Lice ever, the 1848 Hungarian Revolution and freedom
fight against the Austrian Empire interrupted
young Ottó’s formal studies. Probably he wasn’t
Herbaceous Vegetation the best of students anyway, because shortly after
the revolution, his father decided that scholastic
Plants with soft, not woody, stems that die back life wasn’t the way for his son, who continued his
annually. (contrast with woody vegetation) career as a locksmith’s apprentice. Later he went to

Vienna and enrolled in the polytechnic college,
but upon the death of his father, once again he
­discontinued his formal studies. This didn’t mean
that his interest in natural history dwindled, rather
the contrary, he begun to educate himself by auto-
didactic methods and soon built up a reputation
as an outstanding naturalist, versed in many
aspects of zoology, botany and ethnography. In
1875 he was appointed as assistant curator of the
Hungarian National Museum’s zoological collec-
tions. By then he was immersed in the research of
spiders, and between 1876 and 1879 he published
his major entomological work: “The Spider Fauna
of Hungary,” in three volumes. In 1877 he founded
a  journal entitled “Natural History Notebooks”
(Természetrajzi Füzetek). He remained the editor
of this journal for 10 years, during which it became
one of the most popular scientific publications of
the country. In 1888 he embarked on a North
European and Sub-Arctic expedition, following
the route of migratory birds. He passed away
on  December 27, 1914. Ottó Herman produced
eminent works in a number of scientific disci-
plines. Beside entomology, he explored the bird
and fish fauna of Hungary, conducted ethnograph-
ical studies, directed palaeontologic and historical
excavations, and initiated the establishment of
nature reserves and the protection of the natural
environment. He produced 14 books and a great
number of scientific articles. He is regarded as
one  of the most outstanding pioneers of natural
history studies in Hungary.
References
Balázs D (ed) (1993) Magyar utazók lexikona. Panoráma.
Budapest, Hungary
Lambrecht K (1920) Herman Ottó. Az utolsó magyar poli-
hisztor élete és kora. Budapest, Hungary
Hermaphrodite
An individual bearing the features or characteris-
tics of both sexes. Although common among some
Herrich-Schäffer, Gottlieb August
H 1793
invertebrates, hermaphroditism does not occur in
insects.
Hermatobatidae
A family of bugs (order Hemiptera). They also are
known as coral treaders.
 Bugs
Herrich-Schäffer, Gottlieb August
Gottlieb Herrich-Schäffer, son of a medical
­doctor, was born in Regensburg, Germany, on
December 17, 1799. He was influenced in his
early interest in natural history by his great uncle.
By 1817 he had formed a collection of Lepi-
doptera and other insects. In 1818 he entered
Universität Würzburg, and obtained a medical
degree in 1821. However, he used his holidays
to  travel to Erlangen in 1919, to Heidelberg in
1820, to Berlin in 1821, and to Munich in 1822
to further his entomological interests with time
spent in the field. In 1824 he began practicing
as a ­medical doctor in public service and, at the
re­tirement of his father from such ­service in
Regensburg, he returned there in 1828 to take
the  vacant position. He occupied this position
until his retirement in 1856. In 1832 he
befriend­ed C.L. Koch, who was a forestry official
for the Regensburg region (and entomologist),
learned to engrave, and provided 960 engraved
plates for a supplement to Panzer’s “Faunae
in­sectorum germaniae initia.” He continued
Hübner’s “Sammlung europäischer Schmetter-
linge.” In 1835 he published the first part of his
own “Nomenclator entomologicus.” In 1843 he
published the first part of his greatest work, “Sys-
tematische Bearbeitung der Schmetterlinge von
Europa,” with Geyer as its illustrator, completed
in 1856 in six volumes. Other works followed,
including works on exotic species such as “Die
Schmetterlinge der Insel Cuba.” He used to lead
insect-collecting expeditions composed of
1794
H Hesperiidae

f­ amily members and friends in the countryside Hessian Fly, Mayetiola destructor
surrounding Regensburg. He died in Regensburg (Say) (Diptera: Cecidomyiidae)
on  April 14, 1874, leaving a son and four
daughters. richard h. shukle
U.S. Department of Agriculture, Agricultural
­Research Service, West Lafayette, IN, USA
Reference
The Hessian fly, Mayetiola destructor, is a destructive
pest of wheat and occurs in most wheat-­growing
Hofmann O (1874) Necrolog. EntomolZtg (Stettin)
35:277–284 areas of the United States (Fig. 29) and most other
production areas of the world. It is thought to be
endemic to the southern Caucasus and southwest
Asia, the center of origin of the genus Triticum L.,
Hesperiidae and to have dispersed to Europe, North Africa and
North America. While chiefly injurious to wheat, at
A family of butterflies (order Lepidoptera). They
times it causes some damage to barley, rye and triti-
commonly are known as skipper butterflies.
cale. It also can be found on wild grasses such as
 Skipper Butterflies
Agropyron repens (L.), Elmus virginicus L., Hordeum
 Butterflies and Moths
pusillum Nutt., and Aegilops sp., which may serve as
hosts when wheat is not available. The insect received
its common name from Americans when it was
Hesperinidae found infesting wheat on Long Island, New York, in
1779, and it was believed Hessian mercenaries serv-
A family of flies (order Diptera). ing with Lord Howe’s army 3 years earlier had
 Flies brought the pest from Europe in bedding straw.

Hessian Fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae), Figure 29  Distribution of the
­Hessian fly within the United States. (from Foster, J. E., P. L. Taylor, and J. E. Araya. 1986. The Hessian
fly. ­Department of Entomology Agricultural Experiment Station Bulletin 502, Purdue University, West
­Lafayette, Indiana.)
 Hessian Fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae)
Interestingly, the insect was not reported in England
until 1886. It has been recorded in Russia from 1847
onward, especially in the Ukraine, and has been
recognized as a serious pest of wheat in Morocco,
Algeria and Tunisia since the early 1900s.
Life History
As with other flies, development in the ­Hessian fly
has four life stages: egg, larva, pupa and adult. Adults
are small dark to black midges about 3 mm long and
live for 2–3 days after they emerge. Females emit a
sex pheromone to attract males with mating taking
place soon after emergence. As in other members of
the Cecidomyiidae, individual Hessian fly females
generally produce all male or all female progenies,
with the sex of a progeny determined by the female.
Shortly after mating, females begin to deposit eggs
in the grooves along the veins on the upper surface
of the leaves of wheat. The eggs are glistening-red,
cylindrical and about 0.5 mm long. They hatch in
3–7 days depending on temperature, and the ­larvae
crawl down the leaves to feed near the crown in seed-
lings or at the nodes during culm elongation. First-
instar larvae also are red but begin to turn white
within a few days. Second-instar larvae are white,
cylindrical and about 4 mm long when mature. The
cuticle of the mature larva hardens and turns a dark
brown forming a puparium, which is commonly
called a “flaxseed” because it resembles the seed of
flax. Pupation occurs within the puparium.
In northern winter wheat areas, there are typi-
cally two generations a year. In the fall, adults
(Fig.  30) that emerge from infested wheat stubble
infest volunteer wheat or early fall-seeded fields.
Larvae in the fall generation are found in the leaf
sheath at, or near, the crown. With the onset of cold
weather, the mature larva forms a protective pupar-
ium in which it over-winters. In the spring, these
larvae pupate within the puparium and adults
emerge and infest wheat plants about the time they
begin to joint. Larvae in the spring ­generation are
found above the nodes (frequently the second node)
under the leaf sheaths. Larvae pass the summer
H 1795
Hessian Fly, Mayetiola destructor (Say) (Diptera:
­Cecidomyiidae), Figure 30  Hessian fly: adult
female. (from Webster, F. M. 1915. The Hessian fly.
USDA Farmers’ Bulletin 640 – author’ s illustration.)
within the puparia in dry wheat stubble. In late sum-
mer or fall, the larvae pupate and adults emerge to
infest volunteer or early-seeded wheat. In southern
winter wheat areas, supplementary broods can
develop either before or after the main fall genera-
tion, or after the main spring generation. In Canada
and the northern spring wheat areas of the United
States, only one annual spring generation occurs.
Injury to Wheat
Damage in wheat is due entirely to feeding by larvae.
In fall infestations, larval feeding results in stunting
(Fig. 31) and development of a dark green color in
infested primary culms or tillers and can lead to the
death of seedling plants. In spring infestations, larval
feeding prevents normal elongation of internodes,
transport of nutrients to developing grain and results
in lodging at the nodes. The mouthparts of larvae
are highly specialized mandibles that are thought
to inject salivary secretions into the plant. These
secretions are believed to contain enzymes that
inhibit plant growth, increase the permeability of
cell membranes, and soften cell walls.
1796
H Hessian Fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae)
Hessian Fly, Mayetiola destructor (Say) (Diptera:
­Cecidomyiidae), Figure 31  Fall-infested wheat
seedling showing typical stunting. A non-infested
tiller is shown on the left for comparison. (from
Foster, J. E., P. L. Taylor, and J. E. Araya. 1986.
The Hessian fly. Department of Entomology
­Agricultural Experiment Station Bulletin 502, Purdue
University, West Lafayette, Indiana – illustrator
G. Safranski.)
While economic threshold levels have not been
established for Hessian fly infestations, ­evaluation of
injury and loss of winter wheat grain yield indicates
significant damage occurs when fall infestations
exceed 5–8% of tillers, or when spring infestations
exceed 13–20% of stems. When 10% of tillers are
infested in the fall or 20% of stems in the spring,
grain yield loss is estimated at about 250 kg/ha. His-
torically, the Hessian fly has caused losses estimated
at $100 million in a single year. Recently, losses in
Georgia were estimated at $4 million in 1986, and at
$28 million in 1989. Damage from Hessian fly also
has been reported in other southern states; in Texas,
losses estimated at $5 million were reported in 1984.
Control
Because many biotic and abiotic factors ­regulate the
abundance of the Hessian fly, when and where eco-
nomic infestations may occur is not easily predict-
able. Thus, control measures for Hessian fly are
primarily preventive rather than remedial. Preven-
tion of infestation is achieved through (i) planting
resistant wheat varieties, (ii) delaying seeding of
winter wheat to escape fall infestation, (iii) destruc-
tion of volunteer wheat. The use of resistant varieties
is the most economically and environmentally
sound method of control, and has been employed
for the past 50 years. Insecticides are seldom used
for control of Hessian fly. Once larvae have entered
the leaf sheath, a systemic insecticide is required,
and research has demonstrated that resistant wheat
varieties are as effective in controlling damage as
application of systemic insecticide at seeding. If
resistant varieties are not seeded, infestations can be
reduced by cultural practices (i.e., delayed planting
and destruction of volunteer wheat). In much of the
United States cultural practices are effective in
reducing infestations in winter wheat. However, in
the southern United States, delayed planting is not as
effective as it is in northern winter wheat areas
because oviposition and development of larvae can
occur throughout the winter.
Hessian fly Resistance in Wheat
Resistance to Hessian fly has been identified in
common and durum wheat, wild wheat and rye.
Resistance in these sources is expressed as larval
antibiosis; larvae die within 3 days and do not injure
the wheat. Resistance generally is controlled by
single genes that are partially to completely domi-
nant. To date, twenty-nine major genes controlling
Hessian fly resistance in wheat, designated H1
through H29, have been described. Other genetic
 Hessian Fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae)
factors for resistance are those derived from Kawvale
and Marquillo wheat. The genetic control of resistance
in Kawvale has not been determined. Several recessive
genes might control the resistance in Marquillo.
However, recent work has suggested the resistance is
­controlled by a ­temperature-sensitive, partially domi-
nant gene designated H18. The molecular/biochemical
basis of resistance in wheat to the Hessian fly has not
been determined for any gene. However, cytological
studies support the hypothesis that a hypersensitive
reaction is the phenotypic basis of resistance in the
Hessian fly/wheat interaction and involves “recogni-
tion” of an avirulence gene product or process.
Hessian fly Biotypes
The existence of life types or “biotypes” of Hessian
fly, capable of infesting resistant wheat, has been
known for many years. These biotypes describe pop-
ulations that vary in host-adapted alleles. Virulence,
the ability of Hessian fly larvae to survive on and
stunt plants, is controlled by recessive genes at single
loci and operates on a gene-for-gene basis with resis-
tance. Presently, sixteen defined biotypes, Great
Plains (GP) and A through O, have been identified
in field populations by their reaction in a differential
set of four wheat genes for resistance H3, H5, H6 and
H7/8. The Great Plains Biotype is avirulent with
respect to each of these genes for resistance, while
Biotype L is virulent on each of these genes.
Management and durability of
resistance
The development of virulent biotypes of the
Hessian fly capable of surviving on and stunting
formerly resistant wheat is the greatest threat to the
durability of resistance. Several factors in the biol-
ogy of the Hessian fly can affect the development
of virulent biotypes. Dispersal by adults is limited;
adults are short-lived (2–3 days) and oviposition
by females begins one to three hours after mating,
limiting time spent in dispersal. These factors
H 1797
restrict local gene flow, which can increase the
probability of recessive mutations for virulence
becoming homozygous in a population. In opposi-
tion to this are the observations that virulent larvae
can affect the expression of resistance in wheat and
allow avirulent larvae that co-infest to survive. Sur-
vival of avirulent larvae would reduce selection
pressure for virulence in populations and slow the
increase in frequency of virulence alleles associ-
ated with selection from resistant cultivars.
The deployment strategy implemented also can
affect the durability of resistance genes to Hessian
fly in wheat. A sequential release program has been
utilized to date. Under this regime, a single gene is
released in cultivars and used until it begins to lose
effectiveness in the field (generally 6–10 years), at
which time new biotype specific resistance genes are
released. Another strategy is to stack or pyramid
multiple resistance genes. This strategy has not been
­utilized to date because of a lack of suitable markers
to monitor incorporation of multiple resistance
genes; however, the development of molecular mark-
ers will allow implementation of this strategy.
 Wheat Pests and their Management
References
Buntin GD (1999) Hessian fly (Diptera: Cecidomyiidae)
injury and loss of winter wheat grain yield and quality.
J Econ Entomol 92:1190–1197
Cartwright WB, Jones ET (1953) The Hessian fly and how losses
from it can be avoided. USDA Farmers’ Bulletin 1627
Gallun RL (1977) Genetic basis of Hessian fly epidemics. Ann
NY Acad Sci 287:223–229
Grover PB, Shukle RH, Foster JE (1989) Interactions of
­Hessian fly (Diptera: Cecidomyiidae) biotypes on
­resistant wheat. Environ Entomol 18:687–690
Painter RH (1951) Insect resistance in crop plants. Macmillan,
New York, NY, 520 pp
Ratcliffe RH, Hatchett JH (1997) Biology and genetics of the
Hessian fly and resistance in wheat. In: Bondari K (ed)
New development in entomology. Research signpost. Sci-
entific Information Guild, Trivanduram, India, pp 47–56
Ratcliffe RH, Cambron SE, Flanders KL, Bosque-Perez NA,
Clement SL, Ohm HW (2000) Biotype composition of
Hessian fly (Diptera: Cecidomyiidae) populations from
the southeastern, midwestern, and northwestern United
States and virulence to resistance genes in wheat. J Econ
Entomol 93:1319–1328
1798
H Heterobathmiidae

Heterobathmiidae Heterogynidae
A family of moths (order Lepidoptera). They com- A family of moths (order Lepidoptera) also known
monly are known as Valdivian archaic moths. as Mediterranean burnet moths.
 Valdivian Archaic Moths  Mediterranean Burnet Moths
 Butterflies and Moths  Butterflies and Moths

Heteroceridae Heteronemiidae
A family of beetles (order Coleoptera). They A family of walkingsticks (order Phasmatodea).
­commonly are known as variegated mud-loving They commonly are known as common walkingsticks.
beetles.  Walkingsticks and Leaf Insects
 Beetles

Heteroptera
Heteroecious Life Cycle
Sometimes considered to be suborder of
A life cycle where there is alternation of host plants
Hemiptera, containing the “true bugs.”
by insects (generally aphids) through the season. In
 Bugs
species with this type of life cycle, typically the
autumn, winter and spring are spent on a woody
(primary) host plant and summer is spent on an
Heterosis
unrelated herbaceous (secondary) plant. (contrast
with autoecious life cycle)
Also known as hybrid vigor.
 Aphids

Heterotroph
Heterogamy
An organism that obtains energy and other mate-
Alternation of bisexual and parthenogenetic rials by eating other organisms. (contrast with
reproduction. autotroph)

Heterogastridae Heterothripidae
A family of bugs (order Hemiptera, suborder A family of thrips (order Thysanoptera).
Pentamorpha).  Thrips
 Bugs

Heterogynaidae
A family of wasps (order Hymenoptera).
 Wasps, Ants, Bees and Sawflies
Heterozygosity
Having a pair of dissimilar alleles at a locus; a m
­ easure
of genetic variation in a population ­estimated by a
single locus or an average over ­several loci.

Heterozygous
A diploid cell or organism that contains two dif-
ferent alleles of a particular gene.
Hewitt, Charles Gordon
Charles G. Hewitt was born near Macclesfield,
England, on February 23, 1885. He obtained a
B.Sc. from Manchester University in 1902, a
M.Sc. in 1903, and then worked at that university
as a lecturer until 1909, when he received a D.Sc.
degree and emigrated to Canada. His move to
Canada was well timed, as the position of
Dominion Entomologist was vacant. Hewitt had
demonstrated considerable potential while in
England, and despite considerable misgivings
about his level of experience, he was named
Dominion Entomologist of Canada in 1909, and
served in this capacity until 1916. Along they
way, he played a key role in passage of the
“Destructive Insect and Pest Act” of 1910, which
allowed Hewitt to add staff in the form of inspec-
tors and field officers, and to establish entomo-
logical field stations at nine points from coast
to coast. He also published an important treatise
on “House flies and how they spread disease”
in 1912, and “The house fly, Musca domestica
Linn.” in 1914, and served as an officer in several
scientific societies. His position was enlarged
to “Dominion Entomologist and Consulting
Zoologist of Canada” in 1916, and he continued
in this capacity until 1920. His interests and
expertise extended well beyond insects, and he
also was active in natural history and ornithology
organizations. He died prematurely at Ottawa,
Canada, on February 29, 1920. In 1921, his book
titled “Conservation of the wild life of Canada”
was published posthumously, which proved to
be his most renowned contribution to science.
Most importantly, during his brief tenure Hewitt
had developed economic entomology from a
small division to an important branch of the
Department of Agriculture.
Hilltopping
H 1799
Hexapoda
A superclass in the phylum Arthropoda contain-
ing animals with three major body segments and
three pairs of walking legs. Hexapoda consists of
the class Insecta, the insects, and the class Entog-
natha, the collembolans, diplurans, and proturans.
Hibernaculum
An overwintering retreat in which an early-instar
larva retreats or diapauses during the winter. It is
usually made of silk, but may consist of other
materials as well.
Hilarimorphid Flies
Members of the family Hilarimorphidae (order
Diptera).
 Flies
Hilarimorphidae
A family of flies (order Diptera). They commonly
are known as hilarimorphid flies.
 Flies
Hilltopping
Jeffrey H. Skevington
Agriculture and Agri-Food Canada, Ottawa, ON,
Canada
When you are small, locating a potential mate can be
a challenge. Insects employ many strategies of mate
finding, from swarms to individual territories to
specific landmark mating strategies. Landmark mat-
ing strategies involve species setting up mating
aggregations over any conspicuous marker. This can
range from a rock to a tuft of grass, a road, a stream
course, a canyon, a bog, an emergent tree (taller than
1800
H Hilltopping
the others) or a ­hilltop. Although this article focuses
on the latter, this phenomenon is essentially a con-
tinuum. The  difference between simple landmarks
and hilltops is that simple ­landmarks typically sup-
port only a single species. However, emergent trees
in rainforests are likely immensely important for
landmark mating species. Unfortunately, few data
are available on this. Hilltops are significant land-
marks in that they support many species, often
hundreds or in rare cases even thousands. Hilltops
range from massive rocky mountain tops over 4,000 m
high to small hummocks in flat country. These hum-
mocks can function as full-fledged hilltops, particu-
larly if they have some conspicuous landmark on
top. A small rise of land in an otherwise flat area in
Western Australia exemplifies this (Fig. 32); a shel-
ter at the top of the rise serves as a major hilltop-
ping site for many species and families of flies, wasps
and butterflies.
Arguably the best hilltop (most diverse) any-
where in the world is in Queensland, Australia.
Mount Moffatt (1,097 m, 25°03’35” S, 148°02’38” E) is
a distinctive, conical hilltop, rising about 350 m above
the surrounding land (Fig. 32). It is somewhat of a
mystery what makes one hilltop better than another.
Despite this, Mount Moffatt gives us some clues.
The height above the surrounding land is
not too intimidating to exclude many species,
while the hilltop is distinctive and visible for
large distances. The summit is focused and not
very large (about 100 m by 20 m), is variably
vegetated (with a rocky open north end, and
shaded south end) and the habitat around the
hilltop is excellent (in Carnarvon National Park).
Further, no other high points compete for hill-
topping insects within a several kilometer radius.
Hilltops covered entirely by tall trees may also
be important but it is difficult for entomologists
to access these habitats.
The diversity of insects using Mount Moffatt as
an aggregation site almost certainly numbers well
over 1,000 species. Over the course of 3 days between
October 10 and 13, 2002 (early in the ­austral spring),
three entomologists collected 104 species of hilltop-
ping Tachinidae (Diptera) on Mount Moffatt. Based
on this, there are almost certainly over 200 species of
this family alone using Mount Moffatt as a hilltop-
ping aggregation site. Pipunculidae (Diptera) diver-
sity is also high on Moffatt and exceeds 50 species
(Fig.  33). No other groups have been inventoried,
but other abundant and diverse groups using Mount
Moffatt include: Diptera: Anthomyiidae, Bombylii-
dae, Calliphoridae, Muscidae, Pipunculidae, Sarco­
phagidae, Syrphidae, Tabanidae and Tachinidae;
Hymenoptera: Ichneumonidae; and Lepidoptera:
Nymphalidae. More information on hilltopping taxa
is included below.
Hilltopping Taxa
The dominant orders of insects represented on
­hilltops are Diptera, Hymenoptera, and Lepidoptera.
Within Diptera, the following families have been
found to have hilltopping species: Acroceridae,
Agromyzidae, Anthomyiidae, Anthomyzidae, Api-
oceridae, Athericidae, Bibionidae, Bombyliidae,
Calliphoridae, Conopidae, Culicidae, Empididae s.l.,
Fanniidae, Ironomyiidae, Lonchaeidae, Muscidae,
Mydidae, Nemestrinidae, Oestridae, Pelecorhynchi-
dae, Phoridae, Pipunculidae, Platypezidae, Rhagionidae,
Sarcophagidae, Scenopinidae, Sepsidae, Simuliidae,
Sphaeroceridae, Stratiomyidae, Syrphidae, Tabanidae,
Tachinidae, and Therevidae (Fig. 33). Most work on
­hilltopping of Lepidoptera has focused on butterflies
in the following families: Hesperiidae, Lycaenidae,
Nymphalidae, Papilionidae, and Pieridae. Within
Hymenoptera, all families of sawflies appear to
hilltop. Braconidae, Formicidae, Ichneumonidae and
some Sphecidae are also important components of
the hilltopping diversity.
Definition of Hilltopping
Hilltopping is a phenomenon apparently restricted
to insects in which males and virgin or multiple-
mating females instinctively seek a topographic
summit to mate. Larval foodplants or hosts are
­typically not present on the hilltop and may be a
 Hilltopping
H 1801

Hilltopping, Figure 32  (above) A shelter at the top of a small rise of land in an otherwise flat area in
Western Australia serves as a major hilltopping site for many species and families of flies, wasps and
butterflies (Denham, Western Australia (25°54’33” S, 113°31’54” E). (below) Arguably the best hilltop in
the world is Mount Moffatt in Queensland, Australia (1,097 m, 25°03’35” S, 148°02’38” E). This d ­ istinctive,
conical hilltop, rising about 350 m above the surrounding land is a model hilltop in terms of shape,
­vegetation and proximity to other hilltops.

considerable distance away. Hilltopping is a wide- scattered or rare. Mate-finding is difficult under
spread mating system among insects and is focused these conditions.
on groups that are rare, parasitic, predaceous on Note that some species that form behavioral
ephemeral prey, or whose larval foodplants are aggregations do so at hilltops (e.g. Forficula lurida
1802
H Hilltopping

Hilltopping, Figure 33  (Continued)

 Hilltopping
H 1803

Fischer (Dermaptera) and some species of Coc-
cinellidae (Coleoptera)). This prelude to communal
hibernation is not considered hilltopping as defined
here as in such cases sex ratios are typically 1:1 and
no mating or feeding activity typically occurs.
Hilltopping Behavior
General
Typically, there is a very high proportion of male
insects on hilltops. This is explained by the persis-
tence of males in such areas (see below) and the
brief visits by females. Arriving females typically
quickly select a mate and leave the summit. Once
they have mated, males return to the summit while
mated females typically never return. Males often
select bare or open areas from which they can see
approaching females best.
Studies of butterflies reared and released in
different sites suggest that hilltopping species are
genetically programmed to seek out hilltops. Once
there, many species seem to be similarly predis-
posed to seek out very specific places on the hill-
top or otherwise partition the hilltop by their
behaviors. For example, three similar species of
Chalcosyrphus (Diptera, Syrphidae) occur on a
hilltop in Rigaud, Quebec, Canada (45°27’59” N,
74°19’35” W). These species partition the hilltop
by their different behaviours. Chalcosyrphus plesia
(Curran) hovers low over rocks near the summit,
C. curvaria (Curran) typically sits on rocks around
the summit and C. vecors (Osten Sacken) perches
on the leaves of Serviceberry (Amalanchier sp.) or
Red Oak (Quercus rubra) near the summit.

Hilltopping, Figure 33  Collage of hilltopping flies (Insecta, Diptera): (a) Cephenemyia phobifer (Clark)
(Oestridae). Voucher #CNCD144 (Canadian National Collection of Insects-CNC, Ottawa). This rarely
observed fly is common on Mount Rigaud (Quebec, Canada) where this photo was taken. Bot flies are
often found on hilltops despite being found rarely elsewhere (unless they are reared from their hosts). (b)
Cylindromyia sp. (Tachinidae). Voucher #CNCD354. Tachinidae form a major component of most
hilltopping insects. This species is common on Mount Rigaud. (c) Eudorylas sp. (Pipunculidae). Voucher in
University of Guelph Insect Collection (no unique number) (photo by S.A. Marshall). Big-headed flies are
often abundant on hilltops. (d) Euthera tentatrix Loew (Tachinidae). Voucher in CNC (no unique
number). This attractive little fly is rarely collected away from hilltops such as Mount Rigaud (where this
one was photographed). (e) Anthrax maculatus Macquart, (Bombyliidae). Voucher in University of
Queensland Insect Collection (no unique number). Common on hilltops in Eastern Australia. This
specimen was p ­ hotographed on Mount Wellington in Tasmania. (f) Nephrocerus acanthostylus
Skevington ­(Pipunculidae). Voucher #JSS16983 (in CNC). This enigmatic genus of big-headed flies is
rarely collected anywhere. Mount Rigaud has hosted three of the six Nearctic species, including the
specimen in the ­photo. Two ­additional species have been found at a nearby hilltop in Gatineau. This
illustrates how ­effective ­hilltop collecting can be to survey many groups of insects, particularly
parasitoids such as Conopidae, Pipunculidae and Tachinidae. (g) Physocephala marginata (Say) (Conopidae).
Voucher #CNCD347. This is another family of flies that can often be found most readily on hilltops.
Photograph taken on Mount Rigaud. (h) Chalcosyrphus curvaria (Curran) (Insecta, Diptera, Syrphidae) is
one of three closely related flower fly ­species that partition hilltops in northeastern North America by
subtle choices in habitat and differences in behavior. Chalcosyrphus curvaria (Curran) typically sits on
rocks around the summit as in this photo taken at Mount Rigaud, Quebec, Canada (45°27’59” N, 74°19’35”
W). Chalcosyrphus plesia (Curran) hovers low over nearby rocks and C. vecors (Osten Sacken) perches on the
leaves of serviceberry (Amalanchier sp.) and red oak (Quercus rubra) in the same vicinity near the summit.
1804
H Hilltopping
Effectiveness of Hilltopping for Males
Seeking Mates
In theory, if virgin females move randomly upslope,
males patrolling or perching near the summit
should have a higher probability of mating success
than those dispersed throughout the vegetation
below. In years of low population density it should
be adaptive if both sexes tend to move uphill
toward landmarks. However, in years of high pop-
ulation density the numerous males that find their
way to the ridge would compete heavily for a few
females since most would be mated on their uphill
flight. Thus the smaller the population, the lower
the chance of virgin females encountering males
before reaching the hilltop and the more adaptive
the behavior becomes for the males that join the
hilltopping aggregation.
Measuring the adaptive advantage of hilltop-
ping is difficult as males on summits likely face
considerable additional pressures. The energy allo-
cated to hilltopping is likely higher than that
required by individuals remaining off the summit.
Predation rates may also be higher as many preda-
tory insects and birds take advantage of the abun-
dance of prey on hilltops. For example, aerial
foraging birds such as swifts and swallows clearly
focus on hilltop feeding and many large predatory
insects, such as dragonflies (Odonata, Anisoptera)
and robber flies (Diptera, Asilidae), also congre-
gate there to feed. However, given that this may be
the only way for some insects to find a mate, it is
clearly beneficial and has been adopted by a wide
variety of insects.
Evidence for Hilltopping
Mark-recapture studies, particularly using butterflies,
have garnered considerable evidence supporting a
hilltopping strategy as defined here. In one study,
about half of the unmated, reared females released
610 m (117 vertical meters) away from a summit,
were found on the hilltop within 1 day. Mated females
did not go to the summit. In fact, an abnormally high
percentage of female butterflies collected on the
summit were virgin when compared to those cap-
tured elsewhere. The percent virginity for non-hilltop
sites was mostly under 10% in contrast to 35–97% of
females at the summit which were virgin upon arrival.
The apparent scarcity of females at hilltops was due
to inconspicuousness of mating pairs and due to the
fact that virgins stayed only long enough to mate and
non-virgins rarely approached the summit. When
arriving at or released on summit, males strongly
adhered to it. After copulating, females showed little
response to topography.
Mark-release of males of the butterfly Papilio
zelicaon Lucas away from the hilltop resulted in
about 33% returning from varying distances (simi-
lar to the rate of recovery of specimens released
directly on the hilltop). These males returned from
multiple directions and one returned from five km
away and flew into a wind to return. When trans-
ported further away, they would go to another closer
hill; however, there is some evidence that they pre-
fer their own hill when two are similar distances
apart (even though both are used by the species).
The return speed for four P. zelicaon recaptured the
same day varied from 14 (no wind) to 42 (strong
wind at right angle to motion) meters per minute.
In one study, butterflies were found to fly
toward the highest point available. Uphill move-
ment was directed by two cues, the highest location
within a distance of 50 m, and the immediate slopes
available to the butterfly. This suggests that the
highest landmark available in an area is likely to
be used for hilltopping, but this is clearly not always
the case. For example, in an area of high rocky hills in
Tasmania, Australia, the best (most diverse) hilltop
was a partly vegetated hill that was hundreds of
meters lower than the many surrounding hills
(Donaghy’s Hill, 42°11’52” S, 145°55’55” E). More
work is clearly needed to understand how insects
choose specific hilltops and navigate to these
summits.
Differences in behavior between males and
virgin females have been hypothesized to represent
asymmetry in the mate-searching strategy between
genders. This may improve the mate-searching

algorithm by increasing an insect’s chances of
meeting the opposite sex of another of its species
along the way. In butterfly studies, virgin females
were found to perform little searching for males
when not on summits whereas males searched for
conspecifics everywhere.
Timing and Weather
Hilltopping activity is highest in the morning on
sunny, warm days with relatively little wind. Peak
flight times are clearly correlated with temperature
and humidity. Early morning activity (before 8 am)
is marked in very hot, dry climates. Even in moder-
ate climates, the intensity and diversity of hilltop-
pers declines rapidly after mid-day. A resurgence of
activity occurs in some species in the evening
although most species do not appear to have daily
bimodal activity patterns. Crepuscular and noctur-
nal species such as Ormia (Diptera, Tachinidae)
and many moths (Lepidoptera) start to appear at
dusk and some may fly all night. Very little research
has been conducted on nocturnal hilltopping spe-
cies. One note describes the observations of some-
one carrying a lantern up a mountain. No moths
were observed except at the very summit where
hundreds of moths of many species were sitting
everywhere. These insects were there when the
observer arrived and were not attracted to his light.
This clearly illustrates the unexplored potential of
studying nocturnal hilltopping patterns.
The effect of clouds on insect behavior is
extremely pronounced on hilltops. Thousands of
insects can be flying one minute in the sun, produc-
ing a loud hum, when suddenly with the passing of
a cloud over the sun, the insects land and the hum-
ming stops. As the cloud passes, it is as if the entire
hill erupts at once as everything takes to the air
simultaneously. The effect of wind and reduced tem-
perature are similar but not as dramatic because of
the slower shifts. As wind speed increases, activity
moves to the leeward side of the hilltop and remains
focused there until the wind reaches too high a
speed and all activity ceases.
Hilltopping
H 1805
Some species are active throughout the entire
period of peak activity while others have clear
arrival and departure times throughout the day.
Research on the diversity of hilltopping taxa thus
necessitates long hours over many days. Individual
species abundance, particularly of parasitoids, also
varies considerably from year to year. Studies that
span several years may give a much more represen-
tative picture of diversity at a particular summit.
Distances Covered
A few mark-recapture studies have attempted to
determine how far hilltopping individuals travel.
Much remains to be learned here, but it appears
that some species travel considerable distances to
suitable hilltops. Two species of ants, Formica sub-
nuda Emery and Leptothorax muscorum (Nylander),
captured at a hilltop in British Columbia, Canada
returned (even against or across the wind) from
distances up to 760 m away (the farthest tested).
Over 20% of the winged ants released returned to
the hilltop from 760 m away.
Minimum distances traveled by hilltopping
butterflies from the nearest areas of their food-
plants were several thousand vertical feet for Pap-
ilio indra kaibabensis in Arizona and 1,700 vertical
meters for Pieris callidice in France.
It is presumed that once animals initiate a hill-
topping movement they may persistently fly upwards
and cover distances of several kilometers or even
tens of kilometers. This remains to be tested.
Site Tenacity
A substantial number of hilltopping males remain
for periods of time on a summit on a given day.
There is some evidence that some males of some
species may spend their entire lives at a particular
summit. Mark-release and re-sighting studies have
shown that males remain at aggregation sites for
life and that there is a shifting of territories from
day to day. Roosting males of several species have
1806
H Hilltopping
been found at or near the summit of hills, also
supporting the notion that they stay there all of
their lives once they have arrived. One marked
Papilio zelicaon male was seen on a summit on
eight different days over a 20-day period. The lon-
gest period recorded for an individual was 29 days
of one P. zelicaon specimen.
Despite the discovery that male insects of
some species spend most of their lives on a partic-
ular hilltop, it seems unlikely that most species are
this sedentary in their behaviors. At night, during
hot parts of the day, and on days with poor weather,
few diurnal insects can be found anywhere near
the summit. Most presumably leave during these
times to feed or take shelter somewhere below.
Multiple day site fidelity has been demonstrated
in many studies but the reduction in returns over
several days always appears to be steep. The fact that
even successful territory holders rarely spend more
than a single day at a site suggests that males rapidly
deplete their energy reserves, experience a very high
rate of mortality and/or regularly disperse to differ-
ent landmark sites. A sex ratio of 2:1 in favor of males
reported for Cuterebra approxima is hypothesized to
offset high male mortality at aggregation sites.
One study conducted more than 20 years after
the initial study found individuals of Cuterebra
austeni occupying the same landmark site. This spe-
cies used the same perching areas for over two
decades. Other unpublished research on tachinid
flies has found that this is the case for hundreds of
species. This supports the notion that these behav-
iors are hardwired by instincts that have evolved
over millennia.
Survey of Hilltopping Insects
Capture of hilltopping insects typically requires a
focused effort at hand netting. This can be difficult
because of the high degree of habitat partitioning
shown by the species. Three systematists concur-
rently focusing on Diptera collecting on Mount
Moffatt had surprisingly little overlap in 3 days of
collecting Tachinidae. This is presumably due to the
different search strategies employed by these
­dipterists. To sample the diversity of a particular
group, an effort should thus be made to vary the col-
lecting technique. Trapping is typically ineffective, at
least for males. Malaise and pan traps positioned on
hilltops catch a very unrepresentative sample of the
species present. Presumably this is because males are
focused on the task of finding a mate and are not
moving far from their chosen sites. Females tend to
be picked up in these Malaise traps, likely because
they are commuting through the area more than
males. Sugaring (spraying a mixture of artificial
honeydew (honey, Coca-ColaTM and water) on the
leaves of plants) is sometimes effective at attracting
the attention of parasitoid flies on hilltops.
Conservation
The contribution of hilltopping to genetic mixing
between populations may be of great ecological
importance. These aggregation sites may be cru-
cial to the maintenance of populations of many
species and their identification could aid in
attempts to protect threatened populations. Con-
versely, efforts to eradicate pests made sparse by
control measures could be focused on hilltops but
the potential effects on other species would have
to be evaluated before attempting such measures.
Very little effort has been focused on conser-
vation of hilltops, even in parks. Microwave tow-
ers and other communication devices are often
installed even in protected areas on hilltops. Criti-
cal evaluation of these sites is needed before their
development proceeds. The loss of any hilltop
could endanger a particular, or even numerous,
species. The loss of a significant hilltop such as
Mount Moffatt that supports over 1,000 species of
hilltopping insects could have tremendous impacts
on the insect fauna of an area.
Integrated pest management often explores
habitat matrices available in agricultural settings
with the intention of increasing habitat for benefi-
cial insects; however, these strategies have not
explored the importance of hilltops. Given the
 Hinton, Howard Everest
H 1807

critical importance of this mating strategy to many Shields O (1967) Hilltopping. J Res Lepid 6:69–178
Wood DM (1987) Tachinidae. In: McAlpine JF, Peterson BV,
parasitoid insects, more attention should be given
Shewell GE, Teskey HJ, Vockeroth JR, Wood DM (eds)
to these structures in the future. Manual of nearctic Diptera. Agriculture Canada Mono-
graph no 28, Research Branch, Agriculture Canada,
Ottawa, Canada, pp 1193–1269
Yeates D, Dodson G (1990) The mating system of a bee fly
References (Diptera: Bombyliidae). I. Non-resource-based hilltop
territoriality and a resource-based alternative. J Insect
Alcock J (1987) Leks and hilltopping in insects. J Nat Hist Behav 3:603–617
21:319–328
Alcock J, Gwynne DT (1991) Evolution of insect mating sys-
tems: the impact of individual selectionist thinking. In
Bailey WJ, Ridsdill-Smith R (eds) Reproductive behav- Himantopteridae
iour of insects: individuals and populations. Chapman
and Hall, London, UK, 356 pp A family of moths (order Lepidoptera). They also
Alcock J, Smith AP (1995) Landmark-defense and scramble
competition mating systems in two Australian tachinid
are known as long-tailed burnet moths.
flies (Diptera). J Kansas Entomol Soc 68:85–94  Butterflies and Moths
Alcock J, Kemp DJ (2004) Long-term stability in the mating
system of the bot fly Cuterebra austeni (Cuterebridae).
J Insect Behav 17:273–280
Baughman JF, Murphy DD, Ehrlich PR (1988) Population Hindgut
structure of a hilltopping butterfly. Oecologia
75:593–600 The third or posterior section of the gut. The
Beall G (1953) Congregation of butterflies at hilltops. Lepid
hindgut receives products from the midgut at the
News 7:41–43
Britton DR, New TR, Jelinek A (1995) Rare Lepidoptera at juncture of the Malpighian tubules, and consists of
Mount Piper, Victoria – The role of a threatened but- the ileum and/or colon, and the rectum (Fig. 34).
terfly community in advancing the understanding of The waste products exit the hindgut from the
insect conservation. J Lepid Soc 49:97–113
Dodson G, Yeates D (1990) The mating system of a bee fly
anus.
(Diptera: Bombyliidae). II. Factors affecting male terri-  Alimentary System
torial and mating success. J Insect Behav 3:619–636  Alimentary Canal and Digestion
Downes JA (1969) The swarming and mating flight of Dip-
tera. Ann Rev Entomol 14:271–298
Ehrlich PR, Wheye D (1986) “Nonadaptive” hilltopping
behavior in male checkerspot butterflies (Euphydryas Hinton, Howard Everest
editha). Am Nat 127:477–483
McAlpine JF, Munroe DD (1968) Swarming of lonchaeid flies
Howard Hinton was born August 24, 1912, in
and other insects, with descriptions of four new species
of Lonchaeidae (Diptera). Can Entomol 100:1154–1178 Mexico of American parents, but attended high
Merz B (2000) Hilltopping von Dipteren in der alpinen Stufe school and university in Berkeley, California. Then
[Hilltopping of Diptera at alpine levels]. Entomol Basil- he entered Cambridge University in 1934 to study
iensia 22:297–302
O’Hara JE, Skevington JH, Hansen DE (2004) A reappraisal of
for a Ph.D. Between 1939 and 1949 he worked as a
tachinid diversity in Carnarvon N.P., Australia, and esti- taxonomist in the Department of Entomology of
mation of the size of the Australian Tachinidae fauna. the British Museum (Natural History) in London.
Tachinid Times 17:5–8 He published copiously, including a 350-page work
Pe’er G, Saltz D, Thulke HH, Motro U (2004) Response to
topography in a hilltopping butterfly and implications (1945) “A monograph of the beetles associated with
for modeling nonrandom dispersal. Anim Behav stored products.” In 1945 he moved to Bristol Uni-
68:825–839 versity as Reader in Entomology, later being given
Poulton EB (1904) A possible explanation of insect swarms on
a professorial chair and then becoming head of the
mountain-tops. Trans Entomol Soc Lond 1904:23–26
Scott JA (1968) Hilltopping as a mating mechanism to aid the Zoology Department. He was elected a fellow of
survival of low density species. J Res Lepid 7:191–204 The Royal Society in 1961 and was president of the
1808
H Hippoboscidae

Foregut Midgut Hindgut

Gastric
Esophagus Crop Ventriculus Pylorus Ileum Rectum
caecum
Pharynx

Buccal Anus
cavity Proventriculus
Mouth Malpighian tubule

Hindgut, Figure 34  Generalized insect alimentary system (adapted from Chapman, The insects: structure
and function).

Royal Entomological Society of London in 1969–
1970. His research covered many areas of insect
anatomy, physiology, biochemistry and behavior.
But he found time to publish (1967) “Mongooses;
their natural history and behavior” and to found
and edit two journals: “Journal of Insect Physiology”
and “Insect Biochemistry.” He died in Bristol on
August 2, 1977. The 3-volume book “Biology of
insect eggs” on which he had been working just
before his death was published in 1981.
References
Anon (1977) H. E. Hinton. Antenna 1:33
Rothschild M (1978) Howard Hinton – the pharate adult.
Antenna 2:34–36
Hippoboscidae
A family of flies (order Diptera). They commonly
are known as louse flies or keds.
 Flies
Hirsutella
saowanit maimala1, drion g. boucias2
1
Department of Agriculture, Ministry of
­Agriculture and Cooperation, Chatuchak,
­Bangkok, Thailand
2
University of Florida, Gainesville, FL, USA
The genus Hirsutella infects a number of different
types of insects as well as mites and nematodes.
Most of the species (about 50) that infect insects
produce synnemata, i.e., structures composed of a
compact group of erect conidiophores. The most
common species, H. thompsonii, is pleomorphic
and has been separated into three morphologically
distinct groups. Hirsutella thompsonii variety syn-
nematosa occurs in the tropics, whereas varieties
vinacea and thompsonii occur in subtropical and
temperate zones, respectively. All three varieties
can produce conidiogenous structures that are sol-
itary, proliferating phialides that generate one or
more globose, verrucose (warty) conidia, or poly-
blastic conidia that generate subglobose to ellipsoi-
dal conidia with smooth walls. In addition, variety
vinacea produces vinaceous (purple) colonies on
agar, and variety thompsonii characteristically pro-
duces gray-green colonies in culture. Under in vitro
conditions (Ridell mounts), different H. thompsonii
isolates have shown extensive variation in the
length of the phialide, the length of interval between
phialides, the rate of linear growth along substrate,
and in the relative degree of mycelial branching.
The in vivo life cycle of H. thompsonii in host
mites takes only 60–72 h. The conidia invade hosts
(e.g., mites) through the integument. It should be
noted that these short-lived conidia germinate within
minutes after being placed in water. Additionally,
these propagules, unlike the hydrophobic conidia of
many Hyphomycetes, are hydrophilic, and they pos-
sess a mucus coat that facilitates adhesion to the mite
cuticle. It is presumed that these conidia produce
penetrant germ tubes that gain ingress through the

Hirsutella, Figure 35  Scanning electron micrograph of the conidiophores of Hirsutella thompsonii (A), and
(B) represent phialides producing solitary conidia, and (C) phialides producing polybastic conidia.
cuticle. Internalized hyphae break up and form
round, multinucleate chlamydospores, which then
germinate and produce aerial mycelia with conidio-
phores; initially, the mycelia may emerge only
through oral, anal, or genital openings. These obser-
vations suggest that the pathogens may be ingested
and grow out from the gut lumen. In vivo-produced
phialides (Fig. 35) are solitary and arise from mites
attached to plant substrates. Conidiogenesis occurs
within 9–24 h after host death and requires at least
98% relative humidity (RH) and optimal temper-
atures (about 25–30°C). In hot, humid weather,
H. thompsonii can cause spectacular natural epizootics
among mite populations (citrus rust, blueberry, coco-
nut, tomato mites, etc.) and is considered to be a key
natural enemy of various mite pests.
In vitro, this fungus displays a simple growth
cycle. Conidia germinate and produce the mycelial
phase that gives rise to conidiophores and/or chla-
mydospores. Shake flask cultures of several strains
of H. thompsonii var. thompsonii (strain HTF-87,
HtF JAB) produce toxic metabolites including the
protein toxins, Hirsutellin A (HtA) and B (HtB) and
the phomalactones, 6-(1-pro­penyl)-5,6-dihydro-5-
hydroxypyran-2-one. The gene encoding for HtA
(GenBank U86836) has been cloned and sequenced.
This gene codes for a precursor of 164 aa which
includes a 34 aa leader sequence that contains both
a signal and a pro-sequence. This monomeric
­cationic protein undergoes post-translational
Hirsutella
H 1809
­ odification, producing a mature 130 aa HtA toxin
m
which has a calculated Mr=14,159 and pI  =  9.21.
HtA has been reported to possess consensus
sequences similar to those found in various micro-
bial exocellular ribonucleases. Crude filtrates of
mycelial cultures containing HtA were found to be
toxic to wax moth, Galleria mellonella, larvae and
Drosophila melanogaster (Meigen) adults via injec-
tion and per os applications, respectively. In addi-
tion, culture filtrates were active against the citrus
rust mite, Phyllocoptruta oleivora, and the common
two-spotted spider mite, Tetranychus urticae (order
Prostigmata).
References
Chandler D, Davidson G, Pell JK, Ball BV, Shaw K, Suderland
KD (2000) Fungal biocontrol of Acari. Biocontrol Sci
Technol 10:357–384
Jaffee BA (2000) Augmentation of soil with the nematopha-
gous fungi Hirsutella rhossiliensi and Arthrobotrys hap-
tyla. Phytopathology 90:498–504
Mazet I, Vey A (1995) Hirsutellin A, a toxic protein produced
in vitro by Hirsutella thompsonii. Microbiology 141:
1343–1348
McCoy CW (1996) Pathogens of eriophyid mites. In: Lindquist
EE, Sabelis MW, Bruin J (eds) Eriophyid mites: their
­biology, natural enemies, and control. Elsevier, Dordrecht,
The Netherlands, pp 481– 490
Samson RA, McCoy CW, O’ Donnell KL (1980) Taxonomy of
the acarine parasite Hirsutella thompsonii. Mycologia
72:359–377
1810
H Hister Beetles
Hister Beetles
Members of the family Histeridae (order
Coleoptera).
 Beetles
Histeridae
A family of beetles (order Coleoptera). They com-
monly are known as predaceous diving beetles.
 Beetles
Histopathology
A study of abnormal microscopic changes in the
tissue structure of an organism.
History and Insects
thomas r. fasulo
University of Florida, Gainesville, FL, USA
It is impossible to list the number of times insects
affected the course of history, mostly because his-
torians and physicians, during much of our his-
tory, were not aware that many of the diseases that
caused such loss of human life were transmitted
by insects. Even today, medical scientists speculate
over the identity and causes of the “plagues” that
devastated armies and civilian populations in our
early history. While scientists feel confident in
attributing some of these plagues to specific dis-
eases, many of which are vectored by insects, they
cannot be certain.
America, the French and Yellow
Fever
In 1803, Thomas Jefferson, president of the United
States, negotiated one of the greatest land deals in
history. He did so by buying from the French
Emperor Napoleon a vast tract of land known as
Louisiana for the then great sum of $15 million.
This transaction is forever remembered in history
as the Louisiana Purchase. At that time, control of
the city of New Orleans was strategically impor-
tant to the growing United States, as the nation
that controlled New Orleans controlled the Mis-
sissippi River. The United States desperately
needed control of the Mississippi so that western
farmers could ship their grain, hogs, cattle and
other produce to market by flatboats that floated
down the river.
Early in Jefferson’s first administration the
Spanish, who then controlled Louisiana, had
closed the port at New Orleans to the United States’
western settlers and caused them great hardship.
Jefferson knew he needed to secure for his country
a quick and dependable way to transport bulky
goods to market cheaply. But France was then the
most powerful country in the world, and there was
no hope of forcing any privileges from it.
So, in 1802, Jefferson sent James Monroe to
France with an offer to buy New Orleans, and if
possible West Florida, for $2 million. Napoleon
countered with an offer to sell all of Louisiana for
$15 million. Jefferson accepted, doubled the size of
the United States, and ensured that there would
eventually be just one nation spanning the center
of the North American continent. History books
teach us that Napoleon’s primary reason for selling
Louisiana was that he needed the money to pay for
his new war against Great Britain, but this is incor-
rect. The real reason Napoleon sold Louisiana had
to do with mosquitoes.
In January 1802, a flotilla of French transports
carrying 35,000 soldiers commanded by Captain-
General Victor-Emmanuel Charles Leclerc, escorted
by 62 warships, arrived at the island of Hispaniola
in the Greater Antilles. (Today it is divided into the
Republic of Haiti in the west and the Dominican
Republic in the east.) The mission of the force
was to reestablish direct control over the colony
of Saint Domingue (now Haiti and then one of
the richest colonies in the world) as the colony’s
leaders were behaving as if it were an independent

republic. The former black slaves put up a spirited
and well-led defense, but they also had help. By
July, more than 10,000 French troops were dead,
mostly due to yellow fever, a disease transmitted
by mosquitoes. Leclerc himself died of the fever.
His successor requested another 35,000 troops as
reinforcements. Estimates vary as to the number
of French troops Napoleon sent to the island, but
the best recent study suggests that the total
exceeded 60,000 men and women. By November,
only 7,000 starving, sickly troops remained.
­Four-fifths of the French had died of yellow fever,
including 18 generals. By mid-March 1803, France
was once again at war with Great Britain. With
his need for troops to fight the British in Europe,
and with little hope of victory over yellow
fever, Napoleon decided that he could not main-
tain a strong military force in the Americas. He
abandoned his idea to secure Saint Domingue
and to reestablish French power on the North
American continent. On April 30, 1803, he com-
pleted the accords that sold Louisiana to the
United States.
Yellow Fever and the Transatlantic
Slave Trade
The victory of yellow fever, and the mosquitoes
that transmitted it, over the French had far greater
consequences. The collapse of French power in the
West Indies, as their other Caribbean colonies fell
to Britain, negated the argument British slave hold-
ers had that abolishing slavery would not diminish
the transatlantic slave trade, but only turn it over
to the French. In 1807 Britain outlawed slavery for
its own flag vessels. During the late Napoleonic
Wars, Britain began boarding the ships of other
nations and freeing slaves. By the early 1820s, the
British had set their sights on slavery’s abolition
around the world. Due to their efforts, the transat-
lantic slave trade came to an end. With the impor-
tation of inexpensive slaves at an end, the value of
slaves in the southern United States grew, and they
became important enough economic assets that
History and Insects
H 1811
their owners would eventually resort to war to
protect them. Additionally, in 1818, the restored
French monarchy under Louis XVIII considered
an attempt to recover Haiti for France by force.
Survivors of the Leclerc expedition, ­relating their
experiences with yellow fever, helped talk the
French government out of it and possibly pre-
vented a war between France and the United
States.
James Monroe, president of the United States
from 1817 to 1825, introduced the Monroe Doc-
trine (although it was not called that until 20 years
after his death) in December 1823. The Monroe
Doctrine basically stated that the United States
would resist any attempts of “colonization by any
European Powers” in the Western Hemisphere.
When the French later seized control of the
­Mexican government during the American Civil
War of the early 1860s, the United States was too
deeply committed to interfere. However, immedi-
ately at the end of that conflict a large American
army was sent to the Texas-Mexican border. The
French understood the gesture and withdrew their
economic and military support of the government,
which soon fell. A French-American War in the
Western Hemisphere during the nineteenth cen-
tury would have been a real possibility, if France
had maintained colonies there as Spain did. In the
late 1890s American public opinion was directed
against Spain’s remaining colonies in the New
World, and in 1898 The Spanish-American War
took place.
Insects, Greece, Rome and the
Mongols
More than once a powerful army or nation was
defeated or reduced in stature due to insect-vec-
tored diseases. For example, many suggest that
Alexander the Great’s death was most likely due to
malaria, which is transmitted by mosquitoes.
When Alexander died, his driving force died with
him and his empire broke up. Who knows what
might have happened had Alexander lived, to
1812
H History and Insects
expand his empire and establish a dynasty that
would have maintained that empire?
Malaria both hurt and aided the ancient
Romans, who dedicated a temple honoring the
fever goddess. They prayed for mercy from the
mysterious illness that appeared every summer
and made life and travel dangerous. That illness
was malaria, and it both helped and harmed the
Romans. Occasionally barbarian hordes would
descend on mighty Rome during its times of
­ebbing power, only to find themselves dying more
rapidly from malaria than from the famous Roman
short sword. Visigoths, Vandals, Ostrogoths and
Huns would temporarily seize control of the Italian
countryside but, lacking immunity to the local
strain of malaria, would soon die by the thousands. As
the poet Godfrey of Viterbo wrote in 1167, “When
unable to defend herself by the sword,/Rome could
defend herself by means of the fever”.
Rome’s legions also suffered when they ven-
tured into newly conquered areas. Perhaps one
reason the Roman Empire built Hadrian’s Wall in
northern England was to keep out the strain of
malaria that killed half of an 80,000-man army
that tried to conquer Scotland. During the Dark
and Middle Ages, malaria strains to which the sol-
diers weren’t immune decimated entire invading
armies. Death from mosquito-transmitted malaria
has even been suggested as one of the reasons the
Mongols decided not to continue their invasion of
Western Europe after their subjection of much
of Russia and the Balkans. Their raids deep into
Western Europe brought back slaves and loot, but
part of that loot was death to many of the raiders
from the European strains of malaria.
Great epidemics are recorded throughout his-
tory as either having laid waste to great empires or
saving them from invading armies. The oldest
recorded epidemic occurred during the Second
Peloponnesian War in Greece. Large armies
camped in Attica, and refugees crowded the cities.
Diseases spread, but which diseases were involved
isn’t certain. The choices seem to be typhus,
bubonic and pneumonic plague, and smallpox,
with typhus having the most supporters. Typhus
is  transmitted by both fleas and human lice, and
has been associated with war for so long that it
is  also known as “war fever.” In any case, Athens
was defeated and the Golden Age of Greece came
to a close.
Bubonic Plague
Diseases, especially bubonic plague transmitted by
fleas, also helped bring about the decline of the
still strong Eastern Roman Empire. Edward Gib-
bon, in his “The Decline and Fall of the Roman
Empire,” states that “during 3 months, five and at
length ten thousand persons died each day at
Constantinople; and many cities of the East were
left vacant, and that in several districts of Italy the
harvest and vintage withered on the ground.” Thus
did insects and disease cooperate with wars and
political corruption to finally bring the Roman
Empire to its knees. The Roman administrators
died, and their talents died with them. Barbarians
laid waste to civilization and its trappings, and the
Dark Ages settled over much of Europe.
William McNeill, in “Plagues and Peoples,”
states that bubonic plague epidemics occurred in
the eastern Mediterranean region several times
during the late Roman empire. Outbreaks of this
disease were a major factor in preventing Justinian,
emperor of the Byzantine empire, from finding the
required resources needed to restore imperial
unity to the region. Furthermore, flea-transmitted
bubonic plague significantly contributed to the
failure of the Byzantine and Persian empires to
offer effective resistance to the Moslem armies
that swarmed out of Arabia in the middle of the
seventh century.
In China, the first descriptions of the bubonic
plague date from the year 610, or two generations after
its first appearance in the eastern Mediterranean.
A series of devastating epidemics began in 762,
and the records relate that entire provinces lost 50%
or more of their populations. These catastrophes
resulted in a weakening of the existing Chinese
empire, major revolts and an influx of barbarian

influence. China took hundreds of years to regain
pre-plague population levels.
Diseases were even used as an early method
of biological warfare that often spread far beyond
the battlefield and had consequences beyond the
immediate goals of the armies that used them. In
1347, when the Mongols used catapults to hurl
decaying corpses of plague victims over the walls
of Kaffa, a city they were besieging on the shores of
the Crimea, they had no idea that the end result
would be a complete transformation of European
social, educational, religious and political infra-
structures. The bodies carried with them what we
know today as the Black Death, a bacterium that
causes plague in both its bubonic and pneumonic
forms. It is initially spread by fleas, then by fleas
and airborne bacteria after an epidemic starts. The
Black Death, or Black Plague, was one of the major
calamities in history. In Europe alone, it caused
25–75 million deaths in <40 years, depending on
the authority quoted. Most revised estimates place
the total closer to the lower estimate than the
higher.
As Genoese traders fled Kaffa and sailed to
Sicily, they carried with them rats and fleas that
carried the plague. When ships docked at Messina,
most aboard were dead or dying. Soon local inhab-
itants also began to die from the plague. Other
ships carried the Black Death to Italy, where the
results were the same. Tens of thousands died in
the major cities, and many smaller towns and vil-
lages lost all their inhabitants. By the spring of
1349 the Black Death had reached as far as Ireland,
and in 1350 it passed through Scandinavia. Along
its way, it caused moral, religious and political
­disintegration, as well as cruelty. For example, in
many European countries, thousands of Jews were
accused of poisoning the wells, tortured and
killed.
In Europe, there were four major bubonic
plague epidemics in the fourteenth century, with
the last three having significantly declining death
rates as the population developed immunity. When
the first onslaught of the plague was over, one out
of every three Europeans was dead. This was the
History and Insects
H 1813
reason the Black Death had such an effect on
Europe at that time. Imagine a society in which one
out of every three workers, craftsmen, teachers,
religious and political leaders, and others died in
such a short period. With people believing that
they were soon to die, law and order broke down in
many areas and chaos reigned for a time. The very
fabric of society was torn apart. The Black Death
established a dividing line between the central
Middle Ages, with medieval culture in full bloom
and at its greatest strength, and the later Middle
Ages. The later period was one of a significantly
reduced population which required hundreds of
years to attain its pre-Black Death density.
The Black Death Changes Europe
The Black Death is often credited with bringing an
end to the feudal system in Europe. However, that
system was already in decline, although the Black
Death hastened its end. Strong city states and
nations were already emerging, but that process
was drawn out as a result of the Black Death.
Urban populations recovered quickly, in some
cases within a few years, as peasants left the coun-
tryside for better opportunities in the cities. As the
other three epidemics occurred, city dwellers
(many of them former peasants) died, only to be
replaced by more peasants who left the land. But
the Black Death killed many in rural areas, and
this left large areas of farmland without sufficient
workers. With a decreased work force, local lords
either lost wealth and influence or simply went
bankrupt, if they didn’t die of the plague them-
selves. By the fifteenth century, feudalism was in
the past. The surviving workers demanded and
received higher wages and more rights, and many
historians believe that this new sense of freedom
and self-worth led to the Peasants’ Revolt of 1381.
This severe population loss is considered the
main or sole reason for the rise of a monetary
economy and increased technological develop-
ment. When debtors died, their creditors could
not collect as the debtors’ families quite often died
1814
H History and Insects
with them. This helped create a monetary econ-
omy as it allowed immediate payment for many
goods or services. Surprisingly, so many deaths
also led to a better standard of living. Fewer people
meant a decreased demand for goods. This led to
price reductions, which allowed people, now earn-
ing higher wages, to purchase items they could not
have afforded before. In addition, the local lords,
with more land to care for than their work force
allowed, sold off parcels to peasants and towns-
people who never dreamed they would have a
chance to own property. The middle class grew in
size and influence. All this happened because fleas
were busily transmitting the plague.
The fourteenth century was not a good time
for Europe, as it had problems before the arrival of
the Black Death. The economy was in decline. The
rise of the Mongol and Ottoman empires disrupted
important trade routes, and areas of Europe with
seafaring economies were already experiencing
some economic depression. In the interior of
Europe, the overall climate was changing, and the
cooler and wetter weather resulted in lower crop
yields even as the population increased. By the
early 1300s, some parts of Europe were already
experiencing famines. Plus, it didn’t help that The
Hundred Years’ War (1337–1453) added war to
plague and famine. In fact, if the Black Plague had
a beneficial effect, it was that the four epidemics
produced periods of peace in The Hundred
Years’ War.
While the breakdown of class relationships
due to the effects of the Black Death led to a mas-
sive restructuring of society, it also had a lasting
effect on art, literature and religious thought. Cer-
tain professions suffered a higher mortality rate
from the Black Death, especially those whose
duties required them to administer to the sick,
such as doctors and clergy. When the priests died
and no one could hear confession, people learned
that they could live without the church forgiving
their sins. The Black Death is a particularly grue-
some disease, and it caused despair throughout
Christianity. First, people wondered if they had
angered God, and many looked to the church for
guidance. Then they wondered why His chosen
servants in the church were not able to do some-
thing to mitigate the disease and why so many of
the clerics died from it. This caused the Catholic
church to lose much of its prestige, breaking down
blind allegiance to that church and setting the
stage for the Protestant Reformation.
Education of the elite suffered as universities
and schools, usually located in urban areas, were
closed or even abandoned. Up to that time, most
books had been written in either Latin or Greek,
which required rigorous study to learn. But the
Black Death killed many educators and left stu-
dents uneducated or undereducated in Latin and
Greek. This led to a temporary decline in univer-
sities and a long-term decline in the knowledge
of classical languages. The result was a drastic
change in the entire educational system as new
professors and students used the language of the
region instead of Latin or Greek. As a conse-
quence, authors wrote books directly in, and
scholars translated directly to, the vernacular.
Without the need to learn Latin or Greek, more
of the common people could achieve some level
of education.
Due to the death of so many priests and
monks, scribes were authorized to make copies of
manuscripts. This not only greatly increased the
number of people copying manuscripts and books
produced, but it also decreased the price of those
books. During this time the printing press was
invented, which allowed for faster and cheaper
duplication of books. Books, and the knowledge
they contained, were no longer the domain of the
wealthy.
The Black Death even became a part of our
literature. A common children’s nursery rhyme
of  that time survives to this day: Ring a-round
the rosy/Pocket full of posies/Ashes, ashes!/We all
fall down! (Ring a-round the rosy – use your
rosary beads to implore God’s help. A pocket full
of posies – posies were used to hide the odor of
rotting bodies which was also thought to cause
the  plague, so doctors used posies to protect
­themselves from infected plague patients. Ashes,

ashes – the church sanctioned burning the dead
when burying them became too time consuming.
We all fall down – dead.) In Shakespeare’s “Romeo
and Juliet,” a friar is sent to inform Romeo that
Juliet’s apparent suicide was a fake. When the friar
stops to help a family infected with the plague, he
is boarded up in the house with them by their
frightened neighbors. When the friar finally
emerges, he is too late to reach Romeo to tell him
the truth, and both lovers kill themselves over the
other’s apparent and real suicides.
After the four great epidemics in Europe, the
Black Death returned often, but usually only for a
short time and even then in smaller areas. How-
ever, Spain suffered three major outbreaks of plague
during the end of the sixteenth and in the middle
seventeenth centuries. More than half a million
died in the first outbreak and probably as many in
the next two. Together the three epidemics count as
one of the significant factors contributing to Spain’s
decline in economic and political power.
As time passed, people learned how to deal
with the plague, and the mortality rate decreased
sharply as the population gained immunity. One
way people avoided the plague was to leave the
infected area if they could afford it. During the
last big outbreak in Europe, which occurred in
London in 1665, one university professor did just
that and retired to his home in the country. This
was fortunate as it gave him time to compose
his thoughts concerning some theories he held. It
was during this unscheduled vacation that Isaac
Newton worked out the mathematics for his theory
of gravity. Popular history tells us that Newton
developed the theory of gravity because an apple
dropped on his head. We see now that it was
because of a flea.
Although the Black Death finally left Europe in
peace, it continued to plague other areas. Egypt also
experienced a major outbreak of the plague at the
same time it first ravaged Europe, in 1347–1349.
Approximately a third of the population died at that
time, and smaller outbreaks continued to plague
that region over the centuries. During the four-
teenth century many areas of China experienced
History and Insects
H 1815
the plague, and records indicate that up to “two
thirds of the population” died. During the nine-
teenth century, most of the Ottoman governors
and administrators in the Turkish-occupied Bal-
kans lived in the cities and towns where disease,
especially bubonic plague, was common. As they
died they were replaced by more administrators
from Turkey, who also did not have an immunity
to local strains of malaria and other diseases.
Meanwhile, the local Christian populations, most
of whom lived in the rural areas and did not con-
vert to Islam, grew in size as the numbers of Turks
declined. Eventually the Christian populations
revolted, and Ottoman rule was overthrown in the
Balkans.
Just because Europe was no longer ravaged by
the Black Death did not mean that it was free from
the effects of other insect-transmitted diseases.
For example, in 1643, Charles I of England had to
give up his attempt to seize London during the
English Civil War because typhus ravaged his
army, and he missed his chance to quell Parlia-
ment’s revolt. Typhus was originally transmitted
from rats to humans by fleas, but later was most
frequently spread by body lice. But the Class
Insecta does not play favorites and Charles got his
revenge, years after being beheaded, when Oliver
Cromwell died from malaria in 1658. However, at
least one historian presents the case that Oliver
Cromwell was actually poisoned by his enemies,
who used his illness from malaria to disguise his
murder. Even if true, a mosquito still contributed
to Cromwell’s death by assisting in the coverup of
the crime.
Insects and Armies
Diseases have always been closely associated with
large gatherings of armed men and have influ-
enced the outcomes of wars and invasions many
times. Even the greatest of military commanders were
often helpless against disease-transmitting insects.
For example, early in his career, insects dealt Napo-
leon a defeat in his Syrian campaign in the Middle
1816
H History and Insects
East as he boldly attacked into what is now Israel
and Palestine, striking northeast from Egypt. Many
of his troops fell ill from bubonic plague, transmit-
ted by fleas. While total casualties to the plague
were not excessive, the mortality rate was almost
92%. The morale of Napoleon’s soldiers suffered
greatly, and some even  committed suicide when
they contracted symptoms.
Napoleon in Russia
Napoleon, defeated in the West Indies by the mos-
quito and in the Middle East by the flea, suffered
another stunning setback due to an insect – the
human body louse. When he assembled his armies
for the invasion of Russia in June 1812, they num-
bered almost a half million men as they left their
staging centers in Germany and Italy. As the armies
converged and moved through Poland, many of
the soldiers appropriated the insect-­infested hov-
els of the peasants. Sleeping on the straw mats, the
soldiers were infested with the body lice living
there, many of which were carrying typhus. Count-
less thousands were left in makeshift hospitals or
by the wayside and never made it to Russia. By
August 25, Napoleon’s main army had lost 105,000
men out of an original 265,000. By September,
typhus, the prominent disease, and dysentery had
ravaged Napoleon’s armies and some corps were
only half their original size. In October, when the
French entered Moscow, Napoleon’s armies totaled
only 100,000 men, an 80% reduction in strength.
By November, pneumonia had joined typhus and
dysentery in ravaging the Grand Army. For exam-
ple, by December only 20 men remained in the
Third Army Corps commanded by Marshal Ney.
When the Grand Army completed its retreat from
Russia, it could muster only about 7,000–10,000
men fit for duty. This was not the last of Napoleon’s
problems with diseases. In 1814 he raised another
army of 500,000 men only to see it reduced to
170,000 by the Battle of Waterloo. Of the 330,000
casualties, two-thirds were caused by disease,
mostly typhus.
But even Napoleon eventually learned how to
use insects to win battles. When England landed
25,000 troops on the Dutch island of Walcheren
during a campaign in the Netherlands, Napoleon
relied on malaria-transmitting mosquitoes to
defeat this threat to his flank. Of course, at that
time, neither he nor anyone else knew that mosqui-
toes were responsible for spreading malaria. The
malaria was there and that was all that mattered.
The Mexican War of 1847
But insects also could help an army win great vic-
tories on the field of battle even without causing
significant casualties to the opposing force. Dur-
ing the Mexican War of 1847, General ­Winfield
Scott, the commander of a small American army,
led his forces to victory after victory over much
larger Mexican forces, which were often in highly
defensive positions. He did this because mos-
quitoes forced him to take risks. In March of that
year, the American army under Scott’s command
seized Vera Cruz on the Mexican coast. Scott
knew that he had to move into the interior
­without waiting for reinforcements, as the malaria
season was soon to begin. In this case, moving
into the interior also meant gaining considerable
altitude and leaving the mosquito-infested coast
behind. Another general might have waited for
reinforcements and watched his army die from
malaria. But Scott chose to do otherwise, leaving
his supply line hanging in the air, and the result
was a brilliant military campaign.
The Crimean War
The lice and typhus that helped defeat ­Napoleon
remained a problem in later wars. During the
Crimean War of the late 1850s, the Allies and the
opposing Russians were sometimes suffering
12,000 casualties a month from diseases, mostly
typhus, which occurred in two major ­outbreaks.
During the war, the French suffered 200,000

casualties out of an army of 309,000. Only 50,000
of the casualties were from wounds received in
battle; the other 150,000 were hospitalized by
disease. Many of these men were sent to the hos-
pitals because they were infected with typhus but
then died from other diseases, like cholera, that
they contracted in the filthy hospitals. These ter-
rible deaths continued until Florence Nightingale
arrived, reorganized the Allied military hospitals
and reduced the death rate from 45 to 2%. It is
unknown to the author how many casualties the
Russians suffered from typhus.
The American Civil War
Lice played no favorites during the American Civil
War, as they infested Federal and Confederate sol-
diers alike. The armies of both sides were heavily
infested as soldiers often wore and slept in the
same clothes for months. Fortunately, the lice did
not have a typhus reservoir to draw upon as they
fed on Blue and Grey combatants. Although two-
thirds of the 622,000 military deaths during the
American Civil War were caused by disease, typhus
was not a significant cause. According to medical
records, only about 820 deaths were attributed to
typhus. Instead, lice were just an irritant that the
soldiers learned to live with. Federals called the
lice “greybacks,” a name they also used for the
Confederates. And both sides helped pass the time
in camp by playing a gambling game with lice. Sol-
diers placed a tin plate issued for meals over a
candle. As the flame heated the center to the point
where it was just uncomfortable for lice, but not
deadly, the men would place bets and release sev-
eral lice in the center of the plate. The first to
scramble over the edge won money, coffee, tobacco
or fewer hours on guard duty for its recent host.
Today, Civil War reenactors often joke that the
true test of authenticity in a reenactor is when he
does not have to buy his lice from the sutlers
(period merchants), but grows his own.
Dysentery was probably the major non-combat
killer of American Civil War soldiers. Of the
History and Insects
H 1817
approximately 410,000 deaths due to disease, it is
estimated that 40,000–100,000 died from dysen-
tery. Dysentery is caused by unclean conditions,
and the American Civil War armies were just as
unclean as any mass army at that time. People,
even medical personnel, had little awareness of
the good effects a little sanitation would bring.
Latrines were ignored just as often as they were
used. Consider the filth that an army of 75,000–
150,000 men, along with tens of thousands of
horses and mules, creates daily. One month after
General Sherman captured Savannah he began to
ask the government to give him another mission
so he could move his troops from their bivouacs,
which were beginning to accumulate filth. He
knew what disease could do to what was then one
of the finest fighting armies in the world. After a
while he stopped just asking and began to beg.
Eventually, the government allowed him to move
his armies north into the Carolinas. Dysentery
was such a constant companion to the Civil War
soldiers that today’s reenactors sometimes tell
spectators their unit is so authentic that when
they camp they assign a man to defecate upstream
from where they draw their drinking water. Other
armies in other wars also fought battles against
General Dysentery, and the combined casualty
lists were enormous.
Certainly humans can spread filth, but filth-
feeding flies are also a great help. Civil War letters
and diaries are filled with tales of soldiers who
fought the flies more often than they fought the
enemy. With sanitary conditions in a poor state,
and open kitchens as the rule, flies who fed on
filth one minute might be feeding on the armies’
rations the next. The flies also fed on the dead
­soldiers and their horses and mules. During the
American Civil War, as with many wars, General
Dysentery had an aide-de-camp, and his name
was Colonel Fly.
Filth-feeding flies weren’t the only members
of that insect order that helped make life miserable
for the Federals and Confederates (Fig.  36). An
examination of medical records revealed that
­during the American Civil War over 1.3 million
1818
H History and Insects
History and Insects, Figure 36  Confederate soldiers from the American Civil War cover themselves with
sand to avoid mosquitoes (courtesy of the Florida State Archives).
cases of malaria, and 10,000 deaths from that
­disease, were reported just in the Union Army.
While medical records for the Confederates are
nowhere near complete, it is known that in the first
2 years of the war alone, more than one-seventh of
all cases of sickness reported by Confederates
­stationed east of the Mississippi River were due
to malaria. Mosquito-transmitted malaria even
helped stop one of the first attempts by Union
forces to capture Vicksburg.
Many Civil War historians maintain that the
Confederacy lost the Civil War in the West, and
not in the campaigns in Virginia. The loss of Vicks-
burg was the beginning of the end for the Confed-
eracy, and an early victory there would have
hastened the end of the war. A shorter war, without
the horrendous casualties suffered in mid-1864
and later, might have resulted in a more lenient
attitude toward the South after the war. This in
turn might have helped avoid so many of the prob-
lems the United States endured over the next 100
years, problems created by a harsh Reconstruction
that the winners forced on the South. An early
victory might also have left President Lincoln alive
at the end of the war. Only he had the political
influence to counter the political clique known as
the Black Republicans, who dictated policy after
the Civil War. We know from his writings that Lin-
coln intended to be very lenient toward those who
had fought against the Federal government. What
the history of the United States might have been
had he lived is something we can dream of only
wistfully.
World War I
Lice may have played a role in allowing the Allies
to eventually win World War I. As a result of the
early battles of the war, when the Austrians invaded
Serbia and were eventually driven out, northern
Serbia was devastated. Then typhus began to
appear in the Serbian armies in November 1914.
There were fewer than 400 doctors in the country,
most of whom caught the disease, and 126 died.
The country was in ruins: it had almost no experi-
enced nurses, relatively few hospitals for the flow
of wounded, inadequate prisoner-of-war camps
and appalling sanitary conditions. In early 1915, a
typhus epidemic flared up that outpaced any other

recorded typhus epidemic. By April, new cases
numbered in the thousands per day. The mortality
rate among infected patients rose from 20% early
in the disease outbreak, to 60–70% at the height of
the epidemic. Hans Zinsser reports, in his book
“Rats, Lice and History,” that 150,000 people died
in 6 months, Austrian prisoners included. Although
Germany mobilized and went to the aid of Austria
on the Serbian front, there was no troop move-
ment. With the Serbian armies smitten by disease,
the Austrian and German armies stayed put. The
last thing their high commands wanted was a
typhus epidemic destroying their armies. For
almost 6 months, the front was quiet. This delayed
the timetable for the German-Austrian advance
all along the Eastern Front, kept the Russians in
the war longer and may possibly have saved the
Middle East for Britain.
During World War II, the Germans would
transfer units swiftly between the Western and
Eastern fronts, as necessity demanded. However,
during World War I this was not done. For some
reason, although lice were just as common in the
trenches of the Western Front as they were on the
Eastern Front, typhus wasn’t. The battlegrounds of
France and Belgium never experienced the typhus
epidemics of the Eastern Front. As a result, German
and Austrian units were transferred only after
they had undergone a lengthy delousing program.
Generals Louse and Typhus robbed the Germans
and Austrians of the military advantage of interior
lines during World War I.
The Russian Revolution
During the period after the Russian Revolution,
from 1917 through 1923, one Russian scientist
estimated that there were 30 million cases of
typhus, resulting in 3 million deaths, in European
Russia alone. How this might have affected the
political turmoil of this period, when Reds and
Whites were locked in battle for ownership of
Mother Russia, is unknown, but interesting to
speculate about. Even Lenin is supposed to have
History and Insects
H 1819
said, “Either socialism will defeat the louse, or the
louse will defeat socialism.”
Lice
Lice have been our constant companions for most
of history. Only in recent times have many come
to believe that just the poor and uneducated are
infected. In Sweden during the Middle Ages, the
mayor of Hurdenburg was even chosen by a louse.
The eligible elders sat around a table with a louse
in the center and spread their beards. The one
whose beard the louse climbed into was mayor for
the next year. Thomas Becket, the murdered Arch-
bishop of Canterbury, suffered even further dis-
tress after his death. As he was laid out for viewing
in all his woolen clothes, his body cooled and the
parasites began to depart. As one chronicler
reports, “The vermin boiled over like water in a
simmering cauldron, and the onlookers burst into
alternate weeping and laughter.” Lice have even
entered our literature. For example, one of Scottish
poet Robert Burns’ more famous works is entitled,
“To A Louse, On Seeing One On A Lady’s Bonnet
At Church.”
And for all those who wondered why for cen-
turies Europeans insisted on shaving off their hair
only to cover their heads with elaborate wigs, I
give you this simple answer – head lice. (Actually,
head lice were probably only part of the reason
for wearing wigs.) Unfortunately, while an inter-
esting attempt at cultural control, because you
could wash or change most wigs easier than hair,
the fad quickly gained a life of its own. Some wigs
were so elaborate that upper-class women had to
sleep sitting up in bed to prevent destroying what
a hairdresser had spent hours creating. Eventually
people lost track of why wigs were initially worn,
and the wigs were just as infested with head lice as
hair once was. In addition to lice there is one
report of a mouse family setting up housekeeping
in one lady’s elaborate hairpiece. And at least one
noblewoman actually had a bird’s nest woven into
her wig. Eventually wigs were slowly replaced by
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H History and Insects
powdered hair. I’ m not sure why this fashion
came about (cheaper than wigs?), but the white
powder (usually flour) might have served as a
mechanical control for lice.
Mosquitoes
During the Spanish-American War of 1898, the
United States suffered far more casualties from
malaria and yellow fever, transmitted by mosqui-
toes, than it did fighting the Spanish. These casual-
ties occurred just as often in the camps around
Tampa, Florida, as they did in Cuba, Puerto Rico
and the Philippines. Approximately 80% of the
American soldiers in Cuba came down with yel-
low fever. Fortunately, the death rate, while high,
was nothing like that suffered by the French in
Hispaniola at the beginning of the eighteenth
century.
Both before and after the Spanish-American
War, mosquitoes played a major role in preventing
the successful completion of the Panama Canal,
first by a French corporation, then by the Americans.
During these attempts, thousands of workers died,
and tens of thousands of investors lost more than
$3 billion in the French attempt alone, an extremely
large sum at the time. It was only after medical
­scientists realized that malaria and yellow fever
were transmitted by mosquitoes and ordered
that housing conditions be improved (with screen-
ing) and containers of standing water (in which
the mosquito larvae developed) be emptied or
destroyed that the workers stopped dying like flies.
The French seemed to have a continual prob-
lem with disease-transmitting mosquitoes. During
World War I, a French general in Macedonia told
his government that he had to disobey his orders to
attack as his army was in the hospital with malaria.
Fortunately for him, his German opponents were
telling the same thing to their government.
The American military would have other
encounters with malaria transmitted by mosqui-
toes. During World War II, American forces in
the southwest Pacific and southeast Asia suffered
500,000 cases of malaria. General Douglas MacArthur
once complained that two-thirds of his army was
either suffering from malaria or recovering from
it and therefore useless to the war effort. It was
only when he consulted with a malaria-prevention
team, and then commanded that their efforts be
given much higher priority, that the tide of battle
against malaria turned and the Army could then
concentrate on defeating the Japanese. The Allies
also had a secret weapon that came into use
during 1943 – DDT. Used in all theaters of war, in
bases far behind the lines and in newly liberated
cities and towns, DDT was instrumental in reduc-
ing louse-borne disease and malaria. And during
the Viet Nam War, the author remembers regularly
forming up his Marine Corps platoon and walking
the line with a Navy corpsman to ensure every
marine swallowed his malaria prevention tablet.
Tsetse Fly
Outside of Africa, most people encounter tsetse fly
only when reading about the African adventures
of European explorers of the nineteenth century.
But for inhabitants of sub-Saharan Africa, tsetse
fly is much more than an interesting insect from
the history books. Nearly 100 years ago, the tsetse
was identified as the vector of sleeping sickness, a
deadly disease that still has a stranglehold on a
vast area larger than the United States. Tsetse fly,
and the disease it transmits, is the reason why the
ungulate herds of the African savanna have sur-
vived to the present day. This is because, without
modern prophylaxis, livestock cannot live in
regions where the tsetse fly flourishes, and signifi-
cant human losses occur as well. As a result, eco-
nomic development or exploitation of this region,
although attempted many times by African and
European concerns, was never successful. The tse-
tse fly has even been viewed as a beneficial insect
in that it protected central Africa and the huge
ungulate herds from those rapacious European
governments of the nineteenth century that
were busy looting the resources of the rest of the

world. According to the World Heath Organiza-
tion, human sleeping sickness threatens more than
60 million people in 36 countries. Without treatment,
the disease is fatal. In the last century, there have
been three major epidemics of sleeping sickness in
Africa. The last one began in 1970 and is still in
progress as of 2002. The tsetse fly may have pro-
tected central Africa from exploitation, but it had
help from other flies. Even before venturing into
central Africa, European powers had difficulty
penetrating the wall created by malaria- and ­yellow
fever-transmitting mosquitoes.
In cattle, sleeping sickness is called Nagana, and
it is found in an area over 6 million square miles
(or 10 million square kilometers) in Africa. It
affects an estimated 46 million cattle in Africa
alone and causes 3 million cattle deaths annually.
Ninety percent of Africa’s livestock consists of
herds in small villages where maintaining healthy
animals can be the difference between subsistence
misery and a tolerable life for herders and their
families. Other species have also suffered indi-
rectly from this disease. Just in the country of
Zimbabwe alone, more than 2,900 tons of DDT
have been used in an attempt to control the tsetse
fly. The effects of DDT on many vertebrate and
beneficial insect species are too well-known to
mention here. Fortunately, scientists developed
new baiting methods that resulted in a dramatic
decrease in pesticide use.
Black Flies
River blindness, or onchocerciasis, made parts of
Africa and Latin America virtually unlivable. This
disease, caused by a filarial worm, is transmitted
by black flies. The worms migrate from the initial
infection under the surface of the skin throughout
the body and eventually to the surface of the eye,
causing intolerable itching and scarring that results
in a progressive loss of vision. Fortunately, medical
science has developed treatments that are
extremely effective. Recently completed multina-
tional control programs coordinated by the World
History and Insects
H 1821
Health Organization have effectively eliminated
this disease from large parts of West and Central
Africa. This has been accomplished by temporar-
ily (for about 20 years) eliminating the immature
vectors from the breeding sites in rivers with
insecticides, while at the same time therapeutically
controlling the disease in humans. New programs
are being mounted with the hope of continuing
these exciting successes.
Mosquitoes, Disease and Society
Mosquito-transmitted malaria and yellow fever
have also influenced economic and social condi-
tions. To appreciate this you must first understand
how the mosquitoes acquire and then transmit the
diseases. You have probably been wondering, while
reading this article, how these diseases can cause
severe casualties among invading armies but leave
the local residents still standing. Here is a very
simplified explanation of a complex process. There
are many strains of malaria and yellow fever, each
of which is endemic to a region. When children
are born in a region they are usually infected with
the local strain. In children, the diseases may pro-
duce mild reactions, often confined to flulike
symptoms. In many cases, their parents are not
even aware the children have been infected. Sadly,
this is not true for all children, as will be related
later. Children who survive infection by the local
strains may acquire partial immunity to those
strains. However, as people travel to a different
region, they encounter different strains to which
they may not be immune. If they are adults, the
new strains now infecting them can be life-
threatening.
A female mosquito must first bite an infected
person to acquire the disease. After a short period
for the disease organism to complete the next por-
tion of its life cycle, the female mosquito is now
ready to pass it on to another person the next time
it feeds. Many of the more common mosquitoes
that transmit malaria and yellow fever have rela-
tively short flight ranges, often less than a mile.
1822
H History and Insects
Therefore, a female mosquito that acquires the dis-
ease organism from an infected person will not
normally travel great distances to spread that
strain to new areas hundreds or thousands of miles
away. (Unfortunately, modern methods of trans-
portation have helped mosquitoes and disease
strains to overcome these natural barriers.) It is
especially important to know that the mosquitoes
that transmit malaria and yellow fever are either
dawn and dusk feeders and/or night feeders. If you
are new to an area where malaria or yellow fever is
common but take care to protect yourself from
dusk to dawn from biting mosquitoes, then the
chances of becoming infected with and dying from
the local strains of malaria or yellow fever are
greatly decreased. However, there are also at least
two species of yellow fever-transmitting mosqui-
toes that also feed during daylight hours.
One of the lessons learned in Panama was
that an area could be created that was relatively
free of mosquito-borne infections. This area could
then be inhabited by foreign troops, workers,
administrators and their families. The French
began to advocate separate communities for
natives and Europeans. British medical scientists
endorsed this concept by insisting that only natives
carried the disease organisms in their blood. As a
result, representatives of these governments in
many colonies issued regulations against Europeans
socializing or living near local populations. Some
called this segregation, others called it apartheid.
In some countries it eventually took on a social
meaning of its own that was no longer related to
disease transmission.
After the American Civil War, the southern
states were devastated and needed capital invest-
ment to return the region to its former prosperity.
Unfortunately, for most regions in the South, capi-
tal investment never arrived in the amounts nec-
essary. As a result, the southern states would not
achieve economic parity with most northern states
until well into the twentieth century. One of the
reasons for the lack of capital investment was very
simple. The listlessness associated with malaria-
infected workers, as well as the ever-present threats
of malaria and yellow fever epidemics, kept many
northern and European investors from risking
their capital in the South. Workers infected with
malaria may not have died, but the recurring ill-
ness contributed to a significant amount of missed
work days and reduced productivity even when at
work. It was estimated that one-third of all work
missed by southern railroad workers was due to
malaria. Other sectors of the economy also
reported reduced productivity during those
months when malaria was most pronounced.
The Continuing Effects of Insects
on Society
Plagues, famine, social upheaval and conflict, and
the insects that cause them, are easy to write about.
Not as easy to understand and weigh are the every-
day situations we live with that are also influenced
or caused by insects. Numerous insect species
continue to have a direct or indirect effect on the
world economy. Here are a very few examples.
First, there is their entertainment value. Just
as with our ancestors, insects are a part of our
literature. Locust swarms, as depicted in Pearl
Buck’s “The Good Earth” which was also made
into a very good movie, are still very much a
threat for farmers in many regions of the world.
“Leiningen Versus The Ants,” an excellent fic-
tional short story of one man’s battle against army
ants, has been adapted for many movies and TV
dramas. Japanese film-makers used immense cat-
erpillars and moths to destroy Tokyo many times.
The 1950s saw Hollywood produce many films in
which insects were threats to humanity, the
author’s personal favorite being “Them,” a movie
about giant ants. Even the least threatening of
insects have been used to frighten theatergoers.
For example, in “Damnation Alley,” hordes of
Madagascar hissing cockroaches were supposed
to have eaten the entire population of Salt Lake
City. The different ways in which insects have
been featured in articles, books, films and TV
shows is beyond counting. Aside from being

entertaining, all of these have provided jobs
and revenue to the countless people who wrote,
directed or otherwise worked on them. And
probably the world’s shortest poem is even
about an insect. Ogden Nash, the poet laureate of
Maryland until his death in 1971, wrote “Fleas,”
which consists entirely of Adam/Had’m.
In a more serious vein, screwworms, the mag-
gots of some fly species, feed on the flesh of living
mammals. In the early part of the twentieth cen-
tury, these pests cost the United States and Central
American cattle industries millions of dollars in
losses each year. A major effort using sterile flies
has eliminated the primary screwworm from the
United States all the way to southern Panama,
where a barrier is maintained by quarantine and
sterile fly releases to prevent reinfestation from
South America. In 1996, the benefits to the various
cattle industries were estimated at: USA – $796
million, Mexico – $292 million, Central America –
$77.9 million. Before the primary screwworm
threat was eliminated from these regions, many
pet owners accepted that their dogs and cats would
only live with them for a few years. Superficial cuts
suffered by these cherished family members often
resulted in a screwworm infestation, and the pets
would wander off to die.
The urban pest-control industry in the United
States provides hundreds of thousands of jobs and
annually generates more than $5 billion in income.
Termites alone are responsible for more than $1
billion in damage each year. The amount of wood
used to replace such damage is considerable. Its
harvest often generates bad feelings between the
timber industry and environmentalists. Not only
are millions of dollars spent on lawsuits by and
against the timber industry, but the amounts spent
on termiticides and new termite-proof construc-
tion practices take their toll on our economy.
In the United States, the major reason for
most urban children missing school is asthma.
And the major cause of asthma for these children
is an allergic reaction to airborne cockroach feces
and body parts. The social cost of these missed
school days cannot be measured.
History and Insects
H 1823
The urban pest-control industry formerly was
the primary industry responsible for flea control
in the United States. This is no longer true as the
veterinary industry now markets several systemic
insecticides to pet owners to the tune of more than
$250 million dollars a year. This does not include
the amount spent to prevent or cure heartworm,
another mosquito-transmitted disease in dogs
and cats.
And let us not forget the red imported fire ant
(RIFA), which has become such a costly nuisance
in the southern United States. As this ant expanded
its range in the middle of the last century, the
United States spent more than $270 million in an
attempt to destroy it with airborne application of
insecticides. That did not work, and it killed a lot
of the native ant species and allowed the most
aggressive RIFA to move into the vacant niches.
Meanwhile, we continue to spend huge amounts
every year in an attempt to control RIFA, which
has an economic impact on the ornamental, agri-
cultural and livestock industries, as well as others.
In 2002, Congress funded a 5-year, multimillion
dollar program to demonstrate RIFA management
in pastures. Fruit pickers demand extra wages to
work in infested citrus groves. And the amount
spent by homeowners on RIFA insecticides, much
of it wasted as they do not understand how to
apply the baits properly, can only be guessed at.
Visit any hardware/garden store or supermar-
ket and count the number of products specifically
designed to control or manage insects. Scan any
garden or home catalog to see the numerous prod-
ucts, many of which work and others of which do
not, for eliminating insects from your home, gar-
den or life. While many people believe that these
problems and products cost our economy more
than they benefit it, other people believe that they
create even more jobs and strengthen the econ-
omy. Insecticides allow us to feed billions, but
they also generate a cost to human health and
the environment. And these problems are world-
wide in scope. Whether you are growing corn in
the American Midwest or tomatoes in a backyard
in Italy, your insect problems are similar to farmers
1824
H History and Insects
in Central Africa fighting whiteflies on cassava or
rice farmers in the Philippines trying to control
leafminers.
Of course, we now live in the twenty-first cen-
tury, and we expect that the great insect-caused
plagues of the past will not occur again because we
have the technology to control them. Tell that to
the government of Brazil, which employs 50,000
people to go into neighborhoods and empty out
water-filled containers that serve as breeding areas
for disease-vectoring mosquitoes. During late
2001 and early 2002 in Brazil, one out of every 10
workers just in the Brazilian state of Rio de Janeiro
missed work due to being ill with dengue fever,
another mosquito-borne disease. The region’s
association of merchants estimated that 30% of
workers had contracted the disease. Dengue fever
is characterized by severe headache, pain behind
the eyes, high fever, backache, pain in the joints
and a severe rash, with convalescence that may
require several weeks.
Four strains of dengue virus have been identi-
fied, each of which produces lifelong immunity
against the infecting virus. However, exposure
to  infection by a second strain of dengue virus
in  an  already immune individual may result in a
more severe form of dengue known as dengue
­hemorrhagic fever (DHF), with accompanying
­dengue shock syndrome. Death is then common,
particularly when the Type 2 virus infects children
who had previously been infected with another
strain. Increased incidence of DHF has been expe-
rienced in the Western Hemisphere in the last 20
years, with outbreaks occurring in the Caribbean
region. About 205,000 cases of dengue fever were
reported annually in Brazil before the recent 2001–
2002 season. In 1998, an epidemic in that country
caused 540,000 cases. Worldwide, dengue is one of
the most important arthropod-transmitted human
viral diseases, and about 50 million 100 million
cases of DHF are reported annually. On many
Caribbean islands, residents are forbidden to have
bird baths in their yards, and government employ-
ees perform the same function as those in Brazil,
searching for and emptying water containers on
private property. Although it has not experienced a
dengue epidemic in decades, Florida began surveil-
lance for dengue fever in 2002 and by October had
reported nine confirmed and nine probable cases,
all incurred from outside the United States. Texas
occasionally reports cases of dengue fever. Cur-
rently, there is no vaccine available for dengue.
Malaria is also a major problem in Brazil, with
600,000 cases reported annually. Worldwide, 10%
of the world’s population suffers from malaria, and
this mosquito-transmitted disease continues to
kill up to 3 million people every year, including
one child every 30 sec (another authority estimates
one child every 12 sec). Today, malaria is so
uncommon in most industrialized countries that
people think it is a tropical disease. This is not true.
Malaria was once common even in Scandinavia,
and, as previously stated, malaria played a role in
defeating the Roman invasion of Scotland and the
Mongol invasion of western Europe. On the North
American continent the Missouri and Mississippi
watersheds were once major reservoirs for the dis-
ease, as was Canada. Malaria epidemics were once
common on the East Coast, and Staten Island in
New York only experienced heavy development
when extensive draining of marshes occurred after
1900. Today, improvements in housing (screening,
air conditioning, etc.) in many countries make the
severe malaria epidemics of the past unlikely.
However, the malaria virus strains and their vec-
tors continue to evolve, and scientists are con-
cerned that the newer, hardier breeds will one day
make us pay greatly for thinking that it “can’t hap-
pen again.”
Encephalitis viruses are spread by mosqui-
toes. Eastern equine encephalitis (EEE) is the
most lethal of the mosquito-borne encephalitides.
Originally isolated from horses, this virus can also
infect humans. The profitable race-horse industry
can suffer large losses with just the death of a few
highly-valued animals. However, when humans
contract the disease, the cost is far worse. Older
victims usually die, and Florida lost several elderly
adults in 2001, when mosquito populations
were low. When children are infected the cost is

­ articularly high, not even counting the suffering
p For example, the Indian subcontinent, like the
among families. It is estimated that survivors of
EEE each cost society $2.8 million in health care
costs, long-term care and lost productivity.
The West Nile virus, which surfaced in New
York in 1999, moved more quickly across the
United States than many scientists estimated. By
late-2002, it was present in more than 40 states. Yet
with all the publicity surrounding this new disease
to the North American continent, most people
have still not changed their cultural behavior. Per-
haps because, unlike dengue and malaria, not
enough people are dying from it. Parents still
take their children for walks at dusk, mostly shirtless,
in areas where West Nile and other mosquito-
transmitted diseases have been confirmed. Bird
baths are a popular item in many backyards and
are seldom emptied and the water changed. Other
water-filled containers, mostly trash, are commonly
found in yards and lots and serve as breeding
grounds for untold billions of mosquitoes. At least
two primary mosquito vectors of yellow fever and
dengue fever breed in these containers, and both
feed during daylight hours. One, the Asian tiger
mosquito, is an introduced species that is now
firmly established in the southern United States
and is increasing its range.
Will Americans change their cultural habits
only when many of them are sick and dying as the
result of mosquito-transmitted diseases? Probably
not. In South and Central America, and on many
Caribbean islands, where governments often have
more forceful methods of ensuring citizen coop-
eration, many residents still are strong believers in
the right of property and resist government intru-
sion into their yards. As a result, people still die.
The World Health Organization reports that
worldwide there are still 2.5–3 million deaths a
year from malaria, after decades and billions of
dollars spent on educational outreach. And do
not think that West Nile virus is a worst-scenario
disease. There are other mosquito-transmitted
diseases in other regions of the world that are just
waiting for the opportunity to immigrate to North
America, Europe and other continents.
History and Insects
H 1825
rest of Asia, has never experienced a yellow fever
epidemic. What might happen if yellow fever were
to be unleashed on this huge population that has
no immunity to the disease? Andrew Spielman
and Michael D’Antonio in “Mosquito,” an excel-
lent book detailing the natural history of this
group, state that “If yellow fever broke out in India,
for example, where vector mosquitoes and their
human hosts are exceptionally abundant, the loss
of human life would be cataclysmic.” If you believe
this to be an exaggeration, then consider that in
the first half of the twentieth century, India had an
average of 75 million people infected with malaria
at any given time and that the death rate from
malaria was 800,000 a year.
Mosquitoes are such a scourge in India that
they sometimes can accomplish what modern
armies cannot. During mid-2002, swarms of
malaria-bearing mosquitoes along India’s border
with Bhutan were able to drive guerilla insurgents
from their jungle hideouts in search of medical
treatment. The mosquitoes accomplished some-
thing that Indian security forces have been unable
to do for years.
Based on the above, it might seem that human-
ity would be better off if all insects were eliminated.
Fortunately, entomologists, as well as scientists in
many other disciplines, and a growing number of
politicians realize that this is not the case. With
well over a million identified species of insects,
only a few hundred species are listed as serious
pests of humans and their crops, animals and
structures. A few thousand more species can be
considered minor pests. But all the rest are consid-
ered directly or indirectly beneficial to humans.
Just in the United States alone, insects annually
pollinate more than $15 billion worth of fruit and
vegetables. This does not include insects that pol-
linate flowering ornamentals which bring so much
joy to our lives and provide employment in a huge
industry. Plus, the insects themselves are respon-
sible for destroying a thousand times more pest
insects than we are able to with our feeble
“advanced technologies.” As a result, governments,
1826
H History of Biological Control of Wheat Stem Sawflies (Hymenoptera: Cephidae)
universities and private concerns are expanding
their funding for research and implementation
of cultural and biological control techniques that
are successfully managing insect and other pest
populations. Without insects and other arthro-
pods killing other insects and pollinating plants,
the human species probably would have died out,
either through disease or lack of food, even before
leaving the trees.
The next time you swat at a mosquito, or flea,
or other pest, try to remember that its ancestors
did more to change human history than anything
your ancestors ever did. For only when we recog-
nize the tremendous influences insects have had
on the history of our civilization can we begin to
appreciate the adjustments in human behavior
that will be necessary to live acceptably with them
in the future.
References
Bray RS (1996) Armies of pestilence: the impact of disease on
history. Barnes and Noble Books, New York, NY
McNeill WH (1998) Plagues and peoples. Doubleday,
New York, NY
Miller GL (1997) Historical natural history: insects and the
Civil War. Am Entomol 43:227–245
Peterson RKD (1995) Insects, disease, and military history:
the Napoleonic campaigns and historical perception.
Am Entomol 41:147–160
Spielman A, Antonio MD (2001) Mosquito: a natural history
of our most persistent and deadly foe. Hyperion, New
York, NY
History of Biological Control of
Wheat Stem Sawflies
(Hymenoptera: Cephidae)
Thomas G. Shanower
USDA-Agricultural Research Service,
­Sidney, MT,  USA
Wheat stem sawflies (Hymenoptera: Cephidae)
are important pests of wheat and other grain crops
in the northern hemisphere. The egg, larval, and
pupal stages are spent inside the wheat stem,
while the adult is mobile but short-lived. Sawfly
larvae feed inside the stem and the damage they cause
reduces the number and weight of the kernels. The
sawfly larva also cuts the stem at the base of the plant
as it constructs a pupation chamber. The weakened
stem breaks and the grain lodges or falls to the
ground, adding to yield losses and making har-
vesting difficult. Yield losses of more than 20%
have been reported for different sawfly species
from North America, Europe, North Africa and Asia.
The value of these losses may reach $100 million
annually.
Management of wheat stem sawflies has
focused on three strategies: the development of
resistant wheat cultivars, relying primarily on
­solid-stems as a resistance mechanism; various
tillage operations to destroy larvae and/or pupae
in the wheat stubble; and biological control. Pesti-
cides are generally ineffective because the larvae
are hidden within the stem and are too costly for a
relatively low value crop such as wheat. Efforts to
develop solid-stemmed wheat cultivars began in
the 1930s. The first solid-stemmed cultivar was
released in 1946, and more than 20 cultivars have
been released since then. Unfortunately the per-
formance of these cultivars has been variable, and
yields in the absence of sawflies are lower than
hollow-stemmed cultivars. Tillage, either in the
spring or fall, has also been somewhat successful
in reducing sawfly populations but has important
limitations. Tillage requires an additional field
operation, increasing wheat production costs and
reducing profitability, and can also increase soil
erosion rates. The third strategy used to manage
sawflies is biological control using exotic parasi-
toids. Three cephids, Cephus cinctus, C. pygmaeus,
and Trachelus tabidus, have been targets of bio-
logical control programs in Canada and the USA.
These efforts have been extensive, though only
partially successful. The remainder of this section
reviews previous biological control efforts against
these pests, beginning with the biology of sawfly
natural enemies (the parasites, predators and
pathogens that attack these pests).
 History of Biological Control of Wheat Stem Sawflies (Hymenoptera: Cephidae)
Sawfly Natural Enemies
The most important sawfly natural enemies are
parasitic Hymenoptera. A small number of patho-
gens and predators are known to attack sawflies
but have relatively little impact on sawfly popula-
tion dynamics compared to parasitoids. More than
40 parasitoids are reported to attack sawflies,
though little is known about most species; fewer
than ten have a significant impact on sawfly popu-
lations. Many species attack other hosts found in
grasses or other parasitoids (these are then called
hyperparasitoids). The most important primary
parasitoids of grass-feeding sawflies are in the
Braconidae, Ichneumonidae, and Eulophidae fam-
ilies. Before discussing sawfly biological control
programs, it is important to understand the ­biology
and ecology of the key natural enemies.
Several species of braconid attack sawflies in
different regions of the world. All of these bra-
conids are larval ectoparasitoids, living and feed-
ing on the outside of the host larva’s body. The
most important are Bracon cephi and Bracon lis-
sogaster that attack Cephus cinctus, and Bracon
terebella and Heterospilus cephi that attack Cephus
pygmaeus. In general, braconid parasitoids have
more impact on sawfly populations in the New
World than in the Old World. This may be due to
the absence of ichneumonid parasitoids in North
America. In other parts of the world, ichneu-
monids dominate sawfly parasitoid complexes.
Parasitism levels by braconids range from 20 to
more than 90% of available hosts in North
­America, while in Europe and the Mediterranean
region a considerably lower percentage, often
<5%, of hosts are parasitized. The biology of the
braconids is similar among species. Females
locate sawfly ­larvae inside the stems from the
vibrations produced by the feeding larva. She
inserts the ovipositor and paralyzes the larva
before laying one or more eggs. Eggs hatch within
a few days and the braconid larvae begin feeding
on the sawfly larva. Depending on the braconid
species, more than one parasitoid may develop
on a host larva. The braconid larvae consume
H 1827
the entire host except for the head capsule and
epidermis. Larval development is completed in
2–3 weeks, and the parasitoid larva usually over-
winters in the stem of the host plant. There may
be one or two generations per year.
More than ten species of Ichneumonidae
attack sawflies, though most are rare or are present
at very low levels. The most important are species
of Collyria. This group is egg-larval endoparasi-
toids, meaning they live and feed from inside the
body of the host. In Europe, Collyria coxator is the
dominant primary parasitoid of the European
wheat stem sawfly (Cephus pygmaeus). Parasitism
levels of 20–76% have been reported from ­England,
Belgium, France, Switzerland, Germany, Russia,
and Bulgaria. Collyria coxator females locate and
oviposit into sawfly eggs inside the plant stem. Up
to eight Collyria eggs and/or larvae have been
found within a single sawfly (known as superpara-
sitism), though it is not known whether these eggs
are from the same or different females. Despite the
presence of more than one Collyria larva, only a
single adult parasitoid will emerge from a parasit-
ized sawfly larva. Unlike the braconid parasitoids,
Collyria does not paralyze the sawfly larva. The
parasitized sawfly larva continues to feed within
the stem throughout the season, and moves to
the  base of the plant and cuts the stem in a
­manner  similar to an unparasitized sawfly larva.
The ­Collyria larva remains within the host body
until spring when it emerges and pupates. The
adult then chews through the plugged end of
the stub and emerges prior to the emergence of the
unparasitized sawfly adults.
The third and least common group of
­primary parasitoids are several species of Eulo-
phidae. The impact of these species is very low,
and only two species (Pediobius spp.) have been
studied in any detail. These two are found
throughout the northern hemisphere, and are
internal larval parasitoids. In addition to attack-
ing sawfly larvae, they also attack other phy-
tophagous Hymenoptera and Diptera found in
grass stems. Parasitism of more than 50% of
sawfly larvae in grasses has been observed, while
1828
H History of Biological Control of Wheat Stem Sawflies (Hymenoptera: Cephidae)
in wheat, ­parasitism levels rarely exceed 5%. The
reasons for the low attack rate in wheat are
unknown.
Previous Biological Control
Programs
Cephus cinctus was the first sawfly targeted for
biological control. This work was initiated in Can-
ada, and used parasitoids collected in the United
Kingdom. Three parasitoids were selected for
introduction into Canada, two from England (Col-
lyria coxator and Pediobius nigritarsus), and one
from the eastern United States (Heterospilus cephi).
The three species were not collected from C. cinc-
tus but rather from the related sawfly species,
Cephus pygmaeus. From 1930 to 1938, when the
program was discontinued due to the onset of
World War II, more than 450,000 C. coxator and
120,000 P. nigritarsus individuals were released at
sites in Saskatchewan and Alberta. Relatively few
H. cephi adults were released, approximately 1,500,
at one site in Alberta, and in only 1 year. None of
the three released parasitoids became permanently
established, though recoveries of C. coxator were
made up to 2 years after release, with parasitism
up to 9% at some sites.
A second biological control attempt against
Cephus cinctus was undertaken in the western
USA between 1952 and 1955. This program also
used parasitoids obtained from C. pygmaeus,
but this time collections were made in France
instead of England. The predominant parasitoid
collected was again the ichneumonid Collyria
coxator, which parasitized 25–90% of the C. pyg-
maeus hosts. A second parasitoid, Bracon tere-
bella, was also collected. Parasitized C. pygmaeus
cocoons were collected and shipped to Moore-
stown, New Jersey, where they were held for adult
emergence. After emerging, parasitoid adults
were flown to release sites in Montana and North
Dakota. Nearly 40,000 C. coxator and 3200 Bra-
con terebella were released over 4 years. Release
sites were sampled for 6 years beginning in 1953.
No recoveries were made of B. terebella in either
Montana or North Dakota. Low numbers of
C. coxator were recovered at some sites, but
never more than 2 years after release. The last
recovery was in 1957.
The third and smallest biological control pro-
gram against Cephus cinctus began in the late
1950s, when it was recognized that parasitoids
collected from England and France might be
poorly adapted to climatic conditions in western
North America. Parasitoids were collected, again
from the related host C. pygmaeus, in the Ukraine,
the Caucasus region of Russia, and from Sweden.
As in the two previous attempts, the most com-
mon parasitoid collected was Collyria coxator.
However, in laboratory studies, Collyria coxator
suffered high mortality in C. cinctus. Nonetheless,
a small quantity (119) of Collyria coxator from
the Ukraine was released in Alberta. There is no
report of establishment.
The biological control efforts directed at
Cephus cinctus are examples of an approach that
has been called “new associations” or neoclassi-
cal biological control. This approach utilizes nat-
ural enemies from a different but closely related
species to the target host. The target host and
natural enemy thus form a new association, and
according to the theory, establish a lower equi-
librium population level for the target host. In
this instance, four parasitoid species, Collyria
coxator, Pediobius nigritarsus, Bracon terebella
and Heterospilus cephi, were released against
C. cinctus in North America. Despite a sustained
effort, particularly for Collyria coxator, none of
these species was successfully established. The
lack of establishment was likely due to the use of
poorly adapted parasitoids. The parasitoids were
collected from a different sawfly species and the
locations where they were collected, predomi-
nantly England and France, are climatically and
agronomically very different from the North
American Great Plains. Host suitability studies
conducted early in the program could have saved
considerable time and effort by identifying the
lack of compatibility between the target host and
 History of Biological Control of Wheat Stem Sawflies (Hymenoptera: Cephidae)
the natural enemies. In hindsight, these studies
could have avoided the needless release of an
inappropriate parasitoid.
In contrast to the program for Cephus cinctus,
biological control of C. pygmaeus was very suc-
cessful. These efforts, carried out from 1935 to
1938 in the USA and 1937–1940 in Canada,
resulted in the successful establishment of a para-
sitoid and a reduction in sawfly population levels.
Because both C. pygmaeus and T. tabidus were
attacked in Europe by many of the same parasitoid
species, the release program in the US was intended
to address both pest species (Trachelus tabidus has
not been reported from Canada). These programs
utilized Collyria coxator which Canada had
arranged to obtain from England for use against
Cephus cinctus in the western Prairie Provinces
(described above). More than 28,000 C. coxator
adults were released in the US, and more than
24,000 C. coxator females were released in Canada.
Collyria coxator was successfully established in
both the US and Canada, and based on the levels
of parasitism observed in the field, it appears to
play an important role in regulating C. pygmaeus
populations. By the mid-1980s, C. coxator parasit-
ized up to 80% of C. pygmaeus larvae in the field.
This program is an example of classical bio-
logical control, where an introduced pest is con-
trolled by collecting natural enemies from the
target pest’s area of origin, and releasing them into
the new habitat. Once established, the introduced
natural enemies can lower the target pest’s popula-
tion to sub- economic levels.
Summary
Wheat stem sawflies are important pests of wheat
in the northern hemisphere, particularly in North
America. Host plant resistance and cultural con-
trol strategies offer partial control, though both
have significant drawbacks. Attempts at biological
control using exotic parasitoids have been variably
successful. For Cephus pygmaeus and Trachelus
tabidus in eastern North America, an ichneumonid
H 1829
parasitoid (Collyria coxator) was successfully
established and contributes to reduced population
levels now observed in these two sawflies. In con-
trast, biological control for Cephus cintus has been
attempted unsuccessfully three times. Several fac-
tors may have contributed to the lack of establish-
ment of natural enemies in the western US and
Canada. A key factor may have been that the para-
sitoids selected for release were collected from a
different sawfly host and perhaps they were unable
to complete development in this new host. A sec-
ond problem may have been that the parasitoids
were not adapted to the colder, drier climate they
were released into. The release protocol may also
have contributed to the lack of establishment.
Adults were transported across the country (Can-
ada or the US) and released immediately into the
field. This may have resulted in injury or death to
the adult parasitoids due to shipping and ­handling,
and poor timing and/or synchronization between
C. cinctus populations and the parasitoids.
References
Beirne BP (1972) The biological control attempt against
the  European wheat stem sawfly, Cephus pygmaeus.
(Hymenoptera: Cephidae), in Ontario. Can Entomol
104:987–990
Filipy FL, Burbutis PP, Fuester RW (1985) Biological control
of  the European wheat stem sawfly in Delaware
(Hymenoptera: Cephidae). Environ Entomol 14:665–668
Morrill WL, Kushnak GD, Gabor JW (1998) Parasitism of
the wheat stem sawfly (Hymenoptera: Cephidae) in
Montana. Biol Control 12:159–163
Salt G (1931) Parasites of the wheat stem sawfly, Cephus
pygmaeus. Linnaeus, in England. Bull Entomol Res
22:479–545
Shanower TG, Hoelmer KA (2002) Wheat stem sawfly bio-
logical control: past and future. J Agr Urban Entomol
21:197–221
Smith RW (1959) Status in Ontario of Collyria calcitrator
(Grav.) (Hymenoptera: Ichneumonidae) and of Pedio-
bius beneficus (Gahan) (Hymenoptera: Eulophidae) as
parasites of the European wheat stem sawfly Cephus
pygmaeus (L.) (Hymenoptera: Cephidae). Can Entomol
91:697–700
Streams FA, Coles LW (1965) Wheat stem sawfly parasites,
Collyria calcitrator and Pediobius nigritarsis, in eastern
United States. J Econ Entomol 58:303–306
1830
H Hobby, Bertram Maurice
Hobby, Bertram Maurice
Maurice Hobby was born in Southampton, England,
on October 23, 1905. As a schoolboy living in
Southampton, he developed an early interest in
insects and lived close enough to the New Forest
that frequent collecting trips there were possible.
In 1925 he became a student at Oxford University,
gaining his first degree in zoology in 1929. As a
graduate student at Oxford, he worked on “Preda-
ceous insects and their prey,” and gained a D.Phil.
degree in 1934, whereupon he was awarded a
Junior Research Fellowship and, the following
year, was appointed assistant to the chairman of
the entomology department (the Hope Depart-
ment of Entomology). Other positions and pro-
motions followed at Oxford University, until he
retired in 1972. But, in 1938 he was appointed
examiner to the Oxford Delegacy of Local Exami-
nations, and later a delegate of that organization,
as well later still as a Delegate of Oxford University
Department for External Studies. Thus, he served
as a teacher and tutor within the university and
outside it. His research was mainly on predatory
insects, especially Asilidae and Empididae, and
included analyses of insect diets and taxonomy. In
1932, while still a student, he took part in an
Oxford University expedition to Sarawak, bring-
ing back large collections of insects, which were
described in part by him and in part by others. In
1937, he married Marcia Prestidge and, the same
year, joined the ­editorial board of the Entomo­
logist’s Monthly Magazine. He and Marcia had a
45-year association with the editing of that jour-
nal, and it was in this capacity that they had con-
tact with so many entomologist authors. He was
the editor, and a very patient one; she handled
much of the correspondence. He was a member of
entomological societies, president of the Society
for British Entomology in 1937, elected an honor-
ary fellow of the Royal Entomological Society of
London in 1982, and elected a fellow of the ­Linnean
Society in 1948, became a fellow of the Zoological
Society of London, and was secretary of The Ento-
mological Club (an exclusive organization) and
secretary of the Ashmolean Natural History
Society. His ability at swimming led not only to his
participation in swimming and water polo teams
but as Examiner for the Royal Life-Saving Society,
Treasurer of the Oxford University Swim­ming
Club, and President of the Oxfordshire Swimming
Association. He also represented Oxford University
as a chess player. He died at his home in Oxford on
July 19, 1983, survived by Marcia and their one son.
References
Rivers C (1984) Dr Bertram Maurice Hobby, 1905–1983.
Antenna 8:17
Smith KGV et al (1983) Dr. Bertram Maurice Hobby,
1905–1983. Entomologist’s Monthly Magazine
119:179–191
Hocking, Brian
Brian Hocking was born in London on September
22, 1914. With a bachelor’s degree and supportive
wife, Jocelyn, he served as an entomologist in the
Indian army in World War II. In 1946, he was
appointed a faculty member of the Department of
Entomology, University of Alberta, and eventually
became professor and head. His interests were in
insect flight, behavior, migration, and physiology,
especially of Diptera of medical importance, the
diseases transmitted by biting flies, and he studied
the biting flies of Churchill, Manitoba, and means
of their control. He published 110 papers and
supervised research of 30 graduate students. He
was president of the Entomological Society of
Alberta in 1967, and was awarded the Gold Medal
of the Entomological Society of Canada in 1973.
He died on May 23, 1974.
Reference
Riegert PW (1989) Entomologists of Alberta. Entomological
Society of Canada, Ottawa, 56 pp

Hodotermitidae
A family of termites (order Isoptera). They com-
monly are known as Old World harvester
termites.
 Termites
Holarctic Realm (Nearctic and
Palearctic Realms)
The holarctic realm is a large zoogeographic
region encompassing most of North America,
Europe, and northern Asia. It often is subdivided
into the Nearctic realm (North America south to
central Mexico) and Palearctic realm (Europe
and Asia except for Southeast Asia), but the fau-
nas are really quite similar, and characterized by
such fauna as vireos, wood warblers, deer, bison
and wolves.
 Zoogeographic Realms
Holcopogonidae
A family of moths (order Lepidoptera).
 Butterflies and Moths
Holdfast
A structure that functions to attach an insect
to a substrate (e.g., the anal tube of some beetle
larvae).
Holidic Diet
A chemically defined diet. A diet in which all the
components added to the diet are precisely known.
There may be chemical reactions that occur after
mixing, however, that change the nature of the
diet. This type of diet is important for determining
Holland, William Jacob
H 1831
the effects of various nutrients on insect biology.
(contrast with meridic and oligidic diets)
Holland, William Jacob
William Holland was born in Bethany, Jamaica, on
August 16, 1848, and his parents returned with
him to the USA in 1851. His father, a pastor of the
Moravian church, was very interested in natural
­history and brought collected items to Ohio, USA,
from Jamaica. In 1858 the family moved again, to
North Carolina, and in 1863, during the American
Civil War, to Pennsylvania. In 1867 he completed
studies at Moravian ­College, and in 1869 entered
Amherst College, Massachusetts, and studied chem-
istry, physics, geology, astronomy, paleontology, and
philosophy. After graduation he became a school
principal. However, in 1871 he entered Princeton
Theological Seminary at his father’s request, in prep-
aration for a church ministry. In 1874 he became
pastor of a church in Pittsburgh, ­Pennsylvania, and
worked in this position until 1891, when he became
Chancellor of the Western University of Pennsylva-
nia. While still ­occupying the last position, he also
became in 1898 Director of the Carnegie Museum
(in Pittsburgh). His great entomological contribu-
tions were his (1898) “The butterfly book…” and
(1903) “The moth book…” which were affordable
illustrated works on the butterflies and moths of the
USA and Canada. His research was on the Lepi-
doptera of equatorial Africa and of Central and
South America. His collection was rich in lepidopter-
ous specimens from those places, Asia, and the USA.
His interests extended also to fossil reptiles. He died
on December 13, 1932.
References
Holland WJ (1929) Biographical sketch. In: Gunder JD (ed)
The Carnegie Museum, Pittsburgh, Pennsylvania. Ento-
mological News 40:205–217, p. 210–217
Mallis A (1971) William Jacob Holland. In: American ento-
mologists. Rutgers University Press, New Brunswick, NJ,
pp 308–315
1832
H Holocyclic Life Cycle
Holocyclic Life Cycle
A life cycle in which a viviparous parthenogenetic
female produces live nymphs without mating dur-
ing some time of the year, but cyclically produces
males and females that mate and produce eggs.
(contrast with anholocyclic life cycle)
Holometabolous Development
A type of development, also known as complete or
complex metamorphosis, characterized by the
occurrence of the egg, larval, pupal, and adult
stages during the life cycle. This is considered to be
a more evolutionarily advanced form of develop-
ment than hemimetabolous development (incom-
plete or simple metamorphosis).
 Metamorphosis
 Hemimetabolous Development
Holoptic
A term used to describe flies in which the com-
pound eyes meet dorsally.
Holotype
The original specimen from which a new species
description is created. Also referred to as the “type”
specimen.
Homeostasis
Stability or steady state in an ecological commu-
nity or in physiological functions.
Homeothermic
An organism capable of maintaining a relatively
constant body temperature, usually elevated above
the ambient temperature. Although some insects
are capable of limited temperature regulation, they
are poikilotherms, not homeotherms.
 Poikilothermic
Homeotic Gene
Genes that determine the identification and
sequence of segments during embryonic develop-
ment in insects.
Homeotic Genes in Coleoptera
susan j. brown1,2, teresa d. shippy1,2, robin e.
denell1,2, richard w. beeman1,2
1
Kansas State University, Manhattan, KS, USA
2
USDA-ARS-GMPRC, Manhattan, KS, USA
Ever since W. Bateson coined the term “homeotic”
in the late nineteenth century to describe arthro-
pod and vertebrate variants in which one body
part is transformed into the likeness of another,
scientists have been intrigued by the underlying
implication that single genes may have large
effects on the morphological development of
individual segments. Indeed, during the last half-
century, eight homeotic genes located in two
­clusters (Antennapedia Complex  =  ANTC and
Bithorax Complex  =  BXC) have been identified
by genetic and molecular analysis in the fruit fly
Drosophila melanogaster. These genes encode
structurally similar transcription factors that reg-
ulate many other genes to control regional iden-
tity along the anterior-to-posterior axis during
embryogenesis in arthropods and virtually all
other animals. A. Sokoloff and others described
several homeotic mutants in the beetle, Tribolium
castaneum, as early as 1960 in the Tribolium
­Information Bulletin. In 1987, R. Beeman deter-
mined that, similar to homeotic mutations in flies,
Tribolium homeotic mutations display colinearity
in that their order on the chromosome reflects the

order of their functional domains along the ante-
rior-to-posterior axis of the insect. In addition, he
demonstrated that they are located in a single
homeotic complex (HOMC) rather than in two
separate clusters, as in flies. This information
allowed M. Akam and others to infer the ancestral
state of the homeotic complex (single and con-
tiguous) by comparing insect and vertebrate
“Hox” genes.
Genetic and molecular analyses have been
exploited to elucidate the structure and function
of homeotic genes in Tribolium. The evolutionary
counterparts (orthologs) of all eight Drosophila
homeotic genes have been cloned and character-
ized. In addition, the region of the Tribolium
HOMC corresponding to the Drosophila ANTC
has been sequenced. The expression patterns of
the Tribolium homeotic genes are similar to those
of their Drosophila orthologs. The genetic tracta-
bility of Tribolium makes it the most useful non-
Drosophilid arthropod for the study of homeotic
genes. Mutations in all but one of the Tribolium
homeotic genes are now available, and compari-
son of mutant phenotypes between Tribolium and
Drosophila orthologs has yielded several impor-
tant insights into how homeotic gene function
may have evolved in these two insect lineages.
Many phenotypic differences may reflect the
highly derived morphology of Drosophila larvae,
which has internalized head structures and no
external limbs.
Perhaps the most striking case of functional
difference is that of the Tribolium gene, maxillo-
pedia (mxp), and its Drosophila counterpart,
proboscipedia (pb). In Drosophila, pb specifies
the identity of the adult maxillary and labial
appendages. In pb mutants, maxillary append-
ages are abnormal, while labial appendages are
transformed to legs. However, pb has no appar-
ent embryonic function, because larvae lacking
pb function are completely normal. In beetles,
mxp is required for proper maxillary and labial
appendage identity in both larvae and adults. In
individuals completely lacking mxp function,
the maxillary and labial palps are transformed to
Homeotic Genes in Coleoptera
H 1833
legs. Individuals with reduced mxp function
survive to adulthood and show the transforma-
tion of the adult maxillary and labial palps to
legs. These observations suggest that an ances-
tral pb gene may have had an embryonic func-
tion that was lost along the lineage ­leading to
Drosophila, and subsequent analysis of  pb
orthologs in other insects supports this
conclusion.
Mutations in the Drosophila Deformed (Dfd)
gene cause abnormal head development in both
embryos and adults (Fig. 37), but do not produce
homeotic transformations. In Tribolium, embryos
lacking function of the corresponding gene, Tribo-
lium castaneum Deformed (TcDfd), the mandibles
are transformed to antennae and the endite lobes
of the maxillary appendages are missing. This phe-
notype suggests that an ancestral Dfd gene had a
homeotic function.
The embryonic phenotype of Drosophila
Sex  combs reduced (Scr) mutants is somewhat
­controversial. The first thoracic segment (T1) is
unambiguously transformed toward second tho-
racic segment (T2) identity, but the effect on the
Wild-type TcDfd Cx
T1
TcDfd Lu R1/Df(HOMC)
TcDfd mxp Df(HOMC)
Homeotic Genes in Coleoptera, Figure 37  Mutations
caused by expression of homeotic genes.
1834
H Homeotic Genes in Coleoptera
labial segment has been interpreted as either a
transformation to maxillary identity, or merely a
loss of labial identity. In adults, weak Scr alleles
cause T1 to T2, as well as labial to maxillary, trans-
formations. In Tribolium, the embryonic mutant
phenotype of Cephalothorax (Cx) (the ortholog of
Scr) is quite different from that of Scr. In larvae
completely lacking Cx, the labial appendages are
transformed to antennae, and there is no segmen-
tal groove between the T1 segment and the head,
such that the T1 segment is cephalized.
Drosophila mutants lacking Antennapedia
(Antp) function die as larvae and display transfor-
mations of all three thoracic segments toward
more anterior segments. The posterior portion of
T1 is transformed toward posterior labial identity,
and T2 and the anterior portion of T3 are trans-
formed toward T1. During adult development,
patches of T2 leg tissue that lack Antp function are
transformed toward antennal identity. However, in
Tribolium larvae lacking function of prothoraxless
(the Antp ortholog), all three legs are transformed
to antennae.
Null mutants of Drosophila Ultrabithorax
(Ubx) affect segments in the posterior thorax
and the anterior abdomen. One notable effect is
the appearance of a rudimentary thoracic limb
(Keilin’s organ) on the first abdominal segment
(A1) of Ubx mutant larvae. Further studies have
demonstrated that Ubx normally represses
expression of Distalless (Dll) in this segment,
and thus prevents Keilin’s organ formation. In
contrast, during embryonic development in
Tribolium, a pair of modified appendages, called
pleuropodia, transiently appears on A1. Ultra-
thorax (Utx), the Tribolium ortholog of Ubx, and
Dll are co-expressed in the pleuropodia. In
mutants lacking Utx, the pleuropodia are trans-
formed toward legs. Thus, in Tribolium, Utx
modifies appendage identity rather than repress-
ing appendage formation.
The Drosophila abdominal-A (abd-A) gene is
expressed throughout most of the abdomen, but
abd-A mutations cause only the anterior abdominal
segments to be transformed. In contrast, Tribolium
mutants lacking function of Abdominal (A) (the
beetle ortholog of abd-A) develop appendages with
mixed leg/pleuropodia identity on A1–A8. Thus,
the Tribolium abd-A ortholog (A) is responsible
for the limbless abdomen in Tribolium.
The Drosophila Abdominal-B (Abd-B) gene
specifies the identity of the posterior abdomen
and the post-abdomen. These functions are per-
formed by two protein isoforms, M (expressed in
the posterior abdomen) and R (expressed in the
post-abdomen). In Tribolium, expression of the
Abd-B ortholog is limited to the post-abdomen,
and mutant effects are confined to this region as
well. Thus, the Tribolium ortholog seems to lack
the M function. This restriction of the Abd-B
domain also accounts for the expanded region
affected by A mutants.
An overarching theme of the Tribolium
homeotic mutant phenotypes is that removal of
all homeotic genes from a segment results in the
transformation of the appendages on that seg-
ment to antennae (diagrammed in the accompa-
nying figure, first thoracic appendage indicated by
arrow). This is dramatically demonstrated by a
deficiency mutation, (Df(HOMC)), that deletes
most of the Tribolium homeotic genes (TcDfd
through A). Embryos homozygous for Df(HOMC)
have antennae on every segment from anterior
head through posterior abdomen. Another defi-
ciency, (LuR1), which removes all HOMC genes
except A and the Abd-B ortholog, as well as genes
outside the HOMC, has been identified. Although,
individuals homozygous for LuR1 die early in
embryogenesis, embryos carrying one copy of
Df(HOMC) and one copy of LuR1 survive to hatch-
ing. These larvae have normal abdomens, but they
also have antennae on all thoracic and head seg-
ments. These results suggest that the default
appendage state in the absence of homeotic genes
is antenna, and that the presence of homeotic
genes is required to specify other appendage
types, either directly or indirectly.
In Drosophila, artificially induced expression
of most homeotic genes transforms antennae
toward legs. This transformation is apparently
 Honey Bee, Apis mellifera (Hymenoptera: Apidae)
H 1835

caused by repression of homothorax (hth), a gene Homology

required for antennal development. In the
absence of hth and homeotic gene expression, the Homology has been defined as “having a common
antennae are transformed toward legs. Thus, it evolutionary origin,” but also is often used to mean
appears that homeotic gene repression of hth “possessing similarity or being matched.” At the
causes transformation of antenna to leg by uncov- genetic level, it means that organisms have two or
ering an even lower default state of appendage more gene products in common.
identity. Limited data from Drosophila suggest
that homeotic genes also may repress antennal
development within their normal expression Homoplasy
domains. The phenotypes of the Tribolium
homeotic mutants demonstrate that this is a con- The resemblance of organisms due to convergent
served, global property of homeotic genes in evolution or parallelism, rather than to common
insects, and that these genes have dual roles: the ancestry.
specification of regional identity and the repres-
sion of antennal identity.
Homoptera
Sometimes considered to be a suborder of
References
Hemiptera, containing the aphids, whiteflies, scale
insects, leafhoppers, and many other small insects
Akam M (1989) Hox and HOM: homologous gene clusters in
insects and vertebrates. Cell 57:347–349 with piercing-sucking mouthparts.
Beeman RW (1987) A homoeotic gene cluster in the red flour  Bugs
beetle. Nature 327:247–249
Beeman RW, Stuart JJ, Brown SJ, Denell RE (1993) Structure
and function of the homeotic gene complex (HOM-C) in
the beetle, Tribolium castaneum. BioEssays 15:439–444 Homotomidae
Beeman RW, Stuart JJ, Haas MS, Denell RE (1989) Genetic
analysis of the homeotic gene complex (HOM-C) in the A family of bugs (order Hemiptera, superfamily
beetle Tribolium castaneum. Dev Biol 133:196–209 Psylloidea).
Brown SJ, Shippy TD, Beeman RW, Denell RE (2002) Tribo-
lium Hox genes repress antennal development in the  Bugs
gnathos and trunk. Mol Phylogenet Evol 24:384–387
Denell RE, Brown SJ, Brown RW (1996) Evolution of the orga-
nization and function of insect homeotic complexes. Homozygous
Semin Cell Dev Biol 7:527–538

Diploid cells or organisms that contain two identi-

cal alleles of a particular gene.
Homologous Chromosomes

Two or more identical chromosomes. Honey Bee, Apis mellifera

(Hymenoptera: Apidae)

Homologous Genes thomas c. webster

Kentucky State University, Frankfort, KY, USA
Two genes from different organisms and therefore
of different sequence that code for the same gene The genus Apis in the family Apidae consists of at
product. least nine species. This genus is distinguished from
1836
H Honey Bee, Apis mellifera (Hymenoptera: Apidae)
other bee genera by a combination of eusocial
behavior, construction of vertical sheets of bees-
wax comb, communication and recruiting by a
dance language, and a barbed stinger in the worker
bee. Unlike some other bees, Apis species do not
construct material to surround their nests.
The honey bee kept by most beekeepers,
Apis mellifera L., is native to Europe, Africa and
Asia west of the Caspian Sea. Approximately 20
subspecies are distributed across this very large
range. The other Apis species are all native to
southern and eastern Asia. Apis cerana, the spe-
cies most closely related to A. mellifera, is also
managed by beekeepers in China and other
Asian countries.
Anatomy
The honey bee conforms to the general Hymen­o­
pteran morphology. The mandibles are developed
for biting and chewing, and the proboscis for
sucking liquids. Like nearly all apids, Apis species
are adapted to collect and consume nectar and
pollen from flowering plants. Branched hairs
facilitate pollen collection and the pollination of
plants they visit.
The queen bee is the primary reproductive
female. Glands that produce pheromones are in
the mandibles, tergites, rectum and tarsi. Her
abdomen is noticeably enlarged when actively lay-
ing eggs, as her ovaries fill most of her abdomen.
She can extend a curved, unbarbed stinger from
the tip of her abdomen.
The drone is designed entirely for reproduc-
tion. He is easily recognized by his robust thorax
and abdomen, and large compound eyes which
touch each other dorsally and cover most his head.
Most of the abdominal cavity is filled with the
reproductive organs.
The worker bee is a female derived from the
queen form. She is usually sterile, although she has
several small ovarioles in each ovary. A few indi-
viduals in a typical colony are inclined to become
“laying” workers with developed ovaries.
The worker is enabled in her many tasks by
several modifications. Many of the youngest work-
ers become nurse bees, with enlarged hypopha-
ryngeal, salivary and mandibular glands which
secrete food for larvae and the adult queen. Eight
wax glands in the underside of the abdomen are
activated when the colony needs to build comb.
The crop is large enough for the bee to carry nearly
its own weight in nectar or honey. A segment of
each hind tarsus, concave and fringed with hairs,
holds the pollen loads of returning forager bees.
These are corbiculae, or pollen baskets. A straight,
barbed stinger at the tip of the abdomen is used
against intruders, including worker bees from
other hives. The barbs keep it embedded in a large
intruder. When the stinging bee breaks away from
the intruding animal, she leaves the venom sac
attached to the stinger where it continues to pulse
and inject venom. The venom contains a potent
mix of proteins and peptides.
Life History
The egg, larval and pupal stages of each bee are
spent inside of a beeswax cell. The cells are built by
worker bees according to the needs of the colony.
In this way, the workers direct much of colony
growth and reproduction (Fig. 38).
Honey Bee, Apis mellifera (Hymenoptera:
­Apidae), Figure 38  Comb with adult bees, brood
cells and stored honey.

The queen cell begins as an inverted, thimble-
shaped cup. Nurse worker bees add a glandular
secretion, royal jelly, to the cup as the larva grows
inside of the queen cell. Worker bees extend the
cell walls as the larva grows. Soon the cell takes on
a peanut shape, hanging down across the face of
comb. The highly nutritious royal jelly allows the
queen to reach adulthood only 16 days after being
laid as an egg. Within a week or two of emerging
from the cell, the queen takes a mating flight,
returns to the hive, and begins to lay eggs. In the
course of one or several flights, she will mate with
15–20 drones. After mating, her ovaries will enlarge
greatly. She may lay as many as 2,000 eggs daily
during late spring. In temperate climates she will
cease oviposition during winter. The queen rarely
lives more than 2 years. When she begins to fail
due to age or disease, the workers rear a replace-
ment queen, a process called supersedure.
Drone cells are hexagonal and horizontal, usu-
ally built along the bottom and edges of the comb.
Drone larvae are fed a mix of glandular secretions
and honey. The drone emerges as an adult after
24 days of development. He is fertile and prepared for
mating flights within several weeks of adult life. He
may fly a kilometer or more to a drone congregation
area where he may mate on the wing with a young
queen. If he is successful in mating, he then falls away
from the queen, leaving his genitalia inside of her,
and dies soon afterward. In temperate climates, any
drones that survive until autumn are evicted from
their hive by worker bees and die soon afterward.
Most of the cells in the hive are worker cells.
They are hexagonal and horizontal like drone cells,
but smaller. Worker larvae, like drones, consume
glandular secretions and honey. A worker bee takes
21 days to mature in the cell, and emerges as an
adult. She then takes on a complex series of tasks
that collectively maintain the colony. A worker lives
1 or 2 months during the very active period of
spring and summer. Workers winter with their
queen in a quiet cluster in temperate regions.
As a highly eusocial insect, the honey bee is
always colonial. The colony will overwinter with as
few as 10,000 workers and a single queen. Colony
Honey Bee, Apis mellifera (Hymenoptera: Apidae)
H 1837
population grows rapidly in spring, often to over
50,000 workers, several thousand drones and a
queen. Colony fission, or swarming, is normal in
spring. One or more queens are reared in prepara-
tion for this event. The original queen leaves the
hive with a fraction of the colony’s workers and
drones to found a new nest. A daughter queen
remains and inherits her mother’s hive. One or two
afterswarms may leave with other daughter queens
and part of the remaining colony population.
In many regions, honey hoarding ends in
early summer. Late summer and autumn are spent
in reduced foraging, nest maintenance and winter
preparations. In a cold winter, brood rearing and
foraging will cease. The colony clusters tightly to
conserve heat. A healthy, populous colony with
sufficient stored honey will survive until spring,
when it begins again to forage and rear brood.
One honey bee subspecies from southern
Africa, A. mellifera scutellata, was brought to ­Brazil
in 1956 and was later released. It reproduced and
spread rapidly, as it is well adapted to the New
World tropics. Over the following four decades
A. m. scutellata displaced most of the European
race bees from northern Argentina through Mexico,
except where beekeepers maintained European
stock. In the United States it is now established in
many southern states. This African, or Africanized,
bee is undesirable in most countries because of its
extreme tendency to sting, swarm and abscond.
However, it is a much better honey producer in the
tropical countries such as Brazil, where beekeep-
ers have learned to manage it effectively.
Another subspecies, A. m. capensis, is notable
for its ability to produce females from unfertilized
eggs. By this trait the colony can create a new
queen long after its original queen has disappeared.
This bee is native to a small area of South Africa.
However, it has recently begun to colonize the
region further north where it interbreeds with,
and replaces, the colonies of A. m. scutellata kept
by South African beekeepers. Apis mellifera capensis
is a poor honey producer, and very difficult to
exclude from hives. This bee has created a crisis in
South African beekeeping.
1838
H Honey Bee, Apis mellifera (Hymenoptera: Apidae)
Behavior and Physiology
During the first days of adult life, the worker bee
cleans cells. Care for larvae follows, including the
production of glandular secretions. Some workers
will secrete beeswax, build comb, circulate air
through the hive by fanning their wings, ripen
honey, guard the hive entrance, and attend the
queen according to the needs of the colony. The
oldest bees forage for nectar, pollen and water. This
progression creates a division of labor based on
the ages of worker bees in the colony: age poly-
ethism. The transition is regulated in part by juve-
nile hormone secreted by the corpora allata.
The honey bee colony has sophisticated
mechanisms which allow it to respond appropri-
ately to needs and opportunities. One is the flexi-
bility in the workers’ division of labor. The
proportion of the workers in a colony which attend
to a task changes quickly when need arises. The
number of nurses, beeswax producers, guard bees,
water collectors, or foragers increases if more lar-
vae are reared, new comb must be constructed,
enemies threaten, the hive overheats, or flora
become available, respectively.
Another mechanism is the communication of
floral food sources among foragers. A fraction of
the forager bees are scouts, searching for new nec-
tar and pollen sources. Successful scouts commu-
nicate the direction, distance and quality of food
sources to other foragers inside the hive. A waggle
dance in a figure-8 pattern gives this information.
Direction is given by the angle of the dance on the
comb, distance by the duration of the waggling,
and quality by its vigor. Recruit workers that attend
the dance are then able to find the flora, even kilo-
meters from the hive.
The discovery and assessment of potential nest
sites by scout bees is comparable to the cooperative
search for forage. This occurs soon after a swarm
has left its original hive. Individual bees search for
cavities and assess them according to size, location,
dryness and other criteria. These scouts communi-
cate the location and desirability of each site to
the others, by waggle dancing on the surface of
the swarm. This assessment and communication
behavior is analogous to that for food sources.
The queen bee’s presence in the colony and
her condition are communicated to the workers
by pheromones. These pheromones inhibit the
construction of queen cells and ovarian develop-
ment in workers. On a queen’s mating flight, they
attract drones.
Worker bee pheromones play critical roles in
colony life. One comes from the Nasanov gland at
the dorsal tip of the abdomen. Workers release
volatile chemicals from this gland to draw others
to a particular site. When a colony must aggregate
at a new nest site, or unite after a disruptive event,
workers releasing Nasanov pheromone attract the
other colony members. A second pheromone is a
volatile mix at the stinger. When a stinger embeds
in a hive intruder and breaks free of the stinging
worker bee, the sting pheromone is released. This
encourages other bees to sting near the same spot.
By this chemical communication, a quick and
painful defense is mounted against the intruder.
Also, worker and queen bees produce pheromones
that allow them to identify kin and nestmates.
The brood is also a source of pheromone that
inhibits the development of worker ovaries and
stimulates workers to forage. Beeswax stimulates
foraging, apparently due to volatile chemicals in
the wax secreted by the workers’ wax glands.
Honey bees, like all eusocial insects, share
food by passing it directly from one individual to
another. This process, trophallaxis, involves the
entire colony. Trophallaxis informs all colony
members of the availability of food and allows the
efficient transfer of food to storage cells or to those
individuals that most require it.
The temperature of the colony is carefully
regulated. During cold weather, the bees coalesce
into a ball with their branched hairs interwoven.
By rapidly contracting their flight muscles the
workers generate enough heat to maintain the core
of the cluster at around 35°C. In hot weather, water
foragers place droplets of water on the cell walls of
the comb. Other workers perch at the hive entrance
and inside the hive where they pull air through the

hive by fanning their wings. This evaporative cool-
ing keeps the hive at the critical 35°C.
Honey and pollen hoarding is behavior adap-
tive for long periods without forage, especially
winters in cold climates. Honey is made from nec-
tar gathered from the nectaries of many types of
flowering plants. A foraging worker bee carries the
nectar from the flower to another worker waiting
in the hive, which in turn deposits the nectar in a
wax cell. Other workers then ripen the nectar into
honey by adding enzymes and evaporating much
of the water by fanning their wings to circulate air
through the hive. Ripe honey contains glucose,
fructose, approximately 18% water, a small amount
of hydrogen peroxide, and is slightly acidic.
Microbes will not grow in ripe honey. Small
amounts of minerals, amino acids and other nutri-
ents vary in honey according to the floral source.
Pollen is also stored in wax cells, with honey and
glandular secretions added.
Honey is primarily an energy source for adult
bees. Pollen contains protein, vitamins, lipids and
minerals to support brood rearing and the queen
when she is laying eggs. Pollen is not directly con-
sumed by larvae and the queen, however. Nurse
bees feed glandular secretions to them after digest-
ing large amounts of pollen.
Genetics and Breeding
Like all hymenopterans and some insects of other
orders, honey bees are haplodiploid. Females have
two sets of 16 chromosomes, while drones have
one set. When the queen is about to lay an egg she
may apply a bit of semen from that stored in her
spermatheca, as the egg passes through her repro-
ductive tract. This egg is thereby fertilized, and in
most cases becomes diploid and a female. Alterna-
tively, the queen may withhold semen from the egg.
It is then laid as a male, retaining its haploidy. The
development of individuals, in this case males,
from unfertilized eggs, is termed parthenogenesis.
The queen’s ability to control the sex of her
offspring is critical to social life. The rearing of
Honey Bee, Apis mellifera (Hymenoptera: Apidae)
H 1839
drones, which are unable to perform the tasks
necessary to colony maintenance, is relegated to
mating season.
Honey bees differ from many hymenopterans
in their sex determination mechanism. A single
locus controls the sex of the individual bee. A dip-
loid bee that is heterozygous at this “sex locus” is
female. A haploid bee has only one allele at the locus
and hence develops as a male, or drone. Most popu-
lations of honey bee colonies include many alleles
for this locus. Consequently, most fertilized eggs will
be heterozygous and female. However, some frac-
tion of the diploids will be homozygous, depending
on the number of alleles in the population and the
degree of inbreeding. These individuals become
“diploid drones.” They are routinely cannibalized as
very young larvae shortly after they hatch.
The queen bee’s habit of mating far from her
hive and with many drones has important implica-
tions for the colony. It promotes outbreeding and
ensures that most of her diploid offspring will be
heterozygous at the sex locus, hence female. Also,
multiple mating creates multiple subfamilies of
workers within the colony. Worker bees with the
same queen mother and drone father share 75% of
their genes by common descent. Workers from the
same mother and different fathers share only 25%
of their genes by common descent. These relation-
ships are the basis of cooperation and rivalry among
colony members. Much of our understanding of
insect sociality is derived from the genetic systems
of honey bees and other social Hymenopterans.
Honey bee breeding has been fairly success-
ful in many respects. Bees can be bred for gentle
behavior, disease and mite resistance, honey pro-
duction, pollen hoarding and color. However,
breeding is complicated by two factors. One is
the bees’ habit of mating in flight, far from the
hive. Attempts to control mating within an
enclosed structure or on isolated islands have
been unpractical or unsuccessful. Complete con-
trol over breeding relies on instrumental insemi-
nation. This technique works well, but is laborious
when practiced for a large scale operation. The
second factor is the production of diploid drones
1840
H Honey Bees
as lines are inbred to select for a desired trait.
Inbred colonies with queens producing many
diploid drone larvae suffer badly. A closed breed-
ing population must be maintained carefully to
retain sufficient sex alleles.
The honey bee genome was sequenced in 2004.
This allows close inspection of the bee’s traits and
also illuminates comparisons with other species.
 Apiculture
 Bees
 Pollination and Flower Visitation
 African Honey Bee
 Cape Honey Bee
References
Rinderer T (ed) (1986) Bee genetics and breeding. Academic
Press, New York, NY
Ruttner F (1988) Biogeography and taxonomy of honeybees.
Springer Verlag, New York, NY
Seeley TD (1985) Honey bee ecology: a study of adaptation in
social life. Princeton University Press, Princeton, NJ
Snodgrass RE (1956) Anatomy of the honey bee. Cornell Uni-
versity Press, Ithaca, NY
Winston ML (1987) The biology of the honey bee. Harvard
University Press, Cambridge, MA
Honey Bees
Members of the family Apidae (order Hymeno­
ptera, superfamily Apoidae).
 Bees
 Wasps, Ants, Bees and Sawflies
 Apiculture (Beekeeping)
 Pollination an Flower Visitation
Honey Bee Sexuality: An
Historical Perspective
cyrus abivardi
Swiss Federal Institute of Technology Zurich,
Zurich, Switzerland
Before honey bees (Apis mellifera L. and other Apis
species) were domesticated, honey was gathered
from wild hives. Rock painting dating from 8,000
to 15,000 years ago depict a fascinating history of
this process. The earliest recorded hives, which
signify domestication, are about 5,000 years old,
and are seen in paintings and drawings on tombs
and other monuments in Egypt. However, the tra-
ditional hives of Iran and Turkey may provide a
clue to the question of the origin of the hives that
were depicted in Ancient Egypt.
Despite a long history of beekeeping, the first
report on the sexuality of the queen honey bee is
relatively new. Although Aristotle (384–322 b.c.)
noted that some authorities referred to the large
ruler bee as the hive’s mother, he found the hypoth-
esis unlikely, as “nature only arms males.” Because
the hive’s ruler has a sting, Aristotle concluded that
it must be the king and the defenseless drones
were, therefore, the females (Fig.  39). Although
erroneous, Aristotle’s word was taken as virtual
law for the next 1,800 years.
It was not until 1609 that Charles Butler (an
English beekeeper) in his famous book titled “The
Feminine Monarchie,” challenged the idea of king
bees. Thereafter, in 1670, a Dutch scientist (Jan
Swammerdam) proved this hypothesis through
dissection, accompanied by his full anatomical
drawings of queen, drone and worker. He died
in  1680, but his work was not published until
1737–1738.
“The Feminine Monarchie” consists of over
230 pages in ten chapters. The introductory chap-
ter (Cap. I.), titled “Of the nature and properties of
Bees, and of their Queene,” contains information
on the benefits and behavior of honey bees in
general and a description of the Queen bee in
particular.
“Among all the creatures which our bountifull
GOD hath [has] made for the use and service of
man, in respect of (1) great profit with small cost (2)
of their ubiquity or beeing [being] in al [all] countries,
and (3) of their continual labour and consenting
order, the Bees are most to be admired. For first with
the provision of a hive and some little care and
attendance, which need be no hinderance to other
busines [business], but rather a delightful recreatio

Honeybee Sexuality: An Historical Perspective,
Figure 39  Three castes of adult honey bees: above,
the queen; below left, the worker, and below right,
the drone.
amid the same, they bring in store of sweete
delicates most holesome both for meate and
medicine…
“For their order it is such that they may wel [well]
be said to have a comon wealth, since al that they do
is in como [community] without any privat [pri-
vate]… They work for al, they watch for al, they fight
for al. In their private quarrels when they are from the
hive or common treasury howsoever you use them
they wil not resist it by any meanes they can get away.
Their dwelling and diet are common to al alike: they
have like commo [common] care both of their wealth
and young ones. And al this under the government of
one Monarch, of whom above al things they have a
principal care & respect, loving, reverencing, and
obeying her in al things.
“If she goe [goes] forth to solace [rescue] her selfe
(as sometime she wil) many of them attend upon her,
Honey Bee Sexuality: An Historical Perspective
H 1841
garding her person before and behind; they which
come forth before her ever now & the returning, and
looking back, and making with all and extraordinarie
noise, as if they spake [speak] the language of the
knight. Marshalls men, & so away they fly together,
and anon [immediately] in like manner they attend
her back againe [again]. This I may say because I have
seene [seen] it: although the Philosopher be of another
minde [mind].
“If by hir [her] voice she bid them goe [go], they
swarme [swarm]; if being abroad she dislike the
weather, or lighting place, they quickly returne [return]
home again; while she cheereth [cheers] thé [them] to
battaile [battle] they fight; when she is silent they
cease, while she is well, they are cheerfull about their
worke [work], if she droope [droops], they faint also; if
she dy [dies], they will never after prosper, but thence-
forth languish till they bee [be] dead too.
“If they have many Princes, as when two fly away
with one swarme, or when two swarmes are hived to
gether [together]: they strike one of them presently,
and sometime they bring her down that evening to the
matle [battle?], where you may find hir covered with a
little heape [heap] of Bees, otherwise the next day they
carie [carry] her forth either dead or deadly wounded.
Likewise if the olde [old] Queene [Queen] bring forth
many Princes (as she may have six or seaven [seven],
yea sometime halfe [half] a skore or more which
superfluitie [superfluity] nature affordeth [affords]
for more suretie [security], in case some miscarrie
[miscarry]) then lest [in order to avoid] the multitude
of rulers should distract the unstable commons into
factions, within two daies [days] after the last swarme,
you shall finde [find] them that remained, dead
before the hive. I have taken eight of thé [them] up
[above-mentioned] together brought out of one hive,
when two were already gon [gone] forth with the
swarmes…….
“The Queene-bee is a Bee of a comely and
stately shape, browne [brown] of colour as other
Bees, but that her belly is more bright: she is longer
thé [than] honi-bee [honey-bee], by one third part,
that is almost an inch long: she is also bigger then
[than] a honi-bee [i.e., the Worker] but not so big
as a drone, although somewhat longer: hir head
1842
H Honey Bee Sexuality: An Historical Perspective
proportionable, but that it is more roud [round] thé
[than] the little Bees, by reason hir fanges [fangs] be
shorter; hir tonge [tongue] not halfe so long as the
little Bees: for whereas they gather with one nectar,
with the other ambrosia, shee [she] hath no need to
use either, beeing to bee [be] maintained, as other
Princes, by the labor of hir subicects [subjects]; hir
wings of the same size with a smal [small] Bee, &
therefore in respect of hir long body, they seeme
[seem] very short, for they reach but to the middle of
hir nether [lower] part: hir legges [legs], proportion-
able, and of the colour of hir belly, but hir two hin-
legges more yellowe: hir nether part so long, and
halfe so long as hir upper part, more picked thé
[than] smale Bees, and without such three whitish
ringes [rings] as other Bees have at the three parti-
tions: the speere [spear] she hath is but little, and
not halfe so long as the other bees: which, like a kings
sword, is borne rather for shewe [show] and author-
itie [authority], then [than] for any other use: For it
belongeth [belongs] to her subiects as well as to fight
for her, as to provide for her.”
The year 1609 a.d. (i.e., the date of publication
of “The Feminine Monarchie”) is considered by
some to mark the beginning of our knowledge
about the sexuality of the ruler of the hive (the
Queen). However, the first person to understand
the sex of the queen probably was Luis Méndez de
Torres in Spain who published a book in 1586
titled “Tractado breve de la cultivacion y cura de las
colmenas.” In this book, he stated clearly that the
leader bee in the hive laid the eggs, from which all
workers, drones and future queens developed. In
addition to de Torres, the knowledge of several
Persian scientists about the queen bee may be
traced back to a much earlier period.
Ikhawân-ul-Safâ (or The Brethren of Purity),
who flourished in a.d. 950–1000 at Basra, com-
posed a series of tracts in Arabic language known
as the “Epistles of the Brethren of Purity,” (also
reported as “Epistles of the Brethren of Sincerity”)
in the latter half of the tenth century (a.d. 950–
1000). The authors of these series were six ency-
clopaedists, with at least three of them from Persia.
Among their work, the “Dispute Between Man
and  the Animals” is of zoological interest. The
­following citation from this work refers to the sex-
uality of the ruler of the hive:
“There, the king of the hasharât [insects], Ja’ sub
[i.e., the bee queen], rules over the wasps, the flies, the
bugs, the mosquitoes, the dung beetles, the spanish
flies, the butterflies and moths and over the locusts,
i.e., over all small animals which fly by wings, have
no feathers, no bones, no soft hair and no fur.”
The above statement, which uses the Arabic
word of Ja’ sub [the bee queen], clearly refers to the
sexuality of the hive ruler. It is also the first attempt
to define an insect. After about three centuries, al-
Qazwini (the author of “Ajâ’ ib al-Makhluqât,”
composed in a.d. 1263) clearly speaks of the role
of the king of honey bees (the queen) in the pro-
duction of a new king. Despite this report, the
sexuality of the queen as well as the definition of
the insect were overlooked by other scholars for
a long time. While the sexuality of the “King”
remained unnamed until publication of “The
Feminine Monarchie” by Charles Butler in 1609
a.d. (or the publication of the book of Luis Méndez
de Torres in 1586 a.d.), the definition of insect was
ignored until the seventeenth century.
Some citations from al-Qazwini (1263) follow:
“Honeybee is a nice, pleasant animal with fine
body structures…”
“A king has been established to give orders. He
has inherited his honour from his ancestors, since it
is the king who produces again a new king.”
“The king is twice as large as the others and
leads the work…”
Although the word “King” used in the former
sentences may indicate the ignorance of al-
Qazwini about the sexuality of the “Queen,” the
­second phrase (i.e.,“since it is the king who ­produces
again a new king”), leaves no doubt on the aware-
ness of this author about the sex of the “Queen.”
References
Abivardi C (2001) Iranian entomology – an introduction,
2  vols. Springer Verlag, Heidelberg, Germany, 1033 pp

Butler C (1609) The feminine monarchie: a treatise concern-
ing bees, and the due ordering of them wherein the
truth, found out by experience and diligent observation,
discovereth the idle and fond conceipts, which many
have written anent this subject. Joseph Barnes, Oxford
(Facsimile reprinted by De Capo Press, Amsterdam
1969), not paginated
Crane E (1983) The archaeology of beekeeping. Duckworth,
London, UK, 360 pp
Crane E (1985) World perspectives in apiculture. Interna-
tional Bee Research Association, London, UK, 184 pp
Gould JL, Gould CG (1995) The honey bee. Scientific American
Library, New York, NY, 239 pp
Honeydew
whitney cranshaw
Colorado State University, Ft. Collins, CO, USA
Honeydew is the liquid excrement produced by
certain insects in the order Hemiptera that feed on
the phloem of plants. It is an often highly concen-
trated, sugary fluid mixed with some nitrogenous
compounds. Honeydew is primarily produced by
aphids, certain “soft scales,” mealybugs, whiteflies,
some leafhoppers and some treehoppers. Honey-
dew is sometimes used to describe the sugary-rich
materials exuded from certain plant structures
and some insect galls.
The sugars found in honeydew are a complex of
simple plant-derived sugars (e.g., glucose, fructose)
and more complex sugars (e.g., melezitose) that have
usually been derived by the insect using enzymes.
Various nitrogenous materials are found in honey-
dew that primarily reflect plant products of the host
plant, but also includes waste products of the insect.
Nitrogenous materials may occur in concentrations
typically ranging 1–2% in honeydew.
Excreted honeydew is sometimes described
as “dripping sap” or “ghost rain,” particularly when
it drops from shade trees. Some honeydew-pro-
ducing insects may excrete several times their
weight per day. It is sticky and can foul surfaces
where it lands, producing significant nuisance
problems. Surfaces on which honeydew persists
also allow growth of sooty mold fungi.
Hoogstraal, Harry
H 1843
The high sugar content of honeydew may also
attract wasps and bees. This can create nuisance
problems with these insects aggregating as they
forage on shade trees, shrubs and ornamental
plants. However, honeydew is sometimes a valued
resource for honey bees, particularly in Europe
where honey dew-derived honey may be described
as “forest honey,” usually collected when normal
nectar sources are limited and honeydew-producing
insects are common.
Honeydew and other exudates of phloem-
feeding hemipterans have also been used as sweet
materials in the human diet, sometimes described as
manna. The most notable description of this material
is from the Biblical chapter Exodus, and likely
involved a honeydew-producing local scale insect
(Trabutina mannipara) associated with tamarisk.
Honey Pot
A container constructed from cerumen by sting-
less bees and bumblebees and used to store
honey.
Hood
A structure found in soft ticks that projects over,
and covers, the mouthparts.
Hoogstraal, Harry
Emmett R. Easton
University of Hawaii, Honolulu, HI, USA
Harry Hoogstraal was born on February 24, 1917
in Chicago, Illinois. As a young man he and his sis-
ter Catherine were taken by their mother to the
Chicago Natural History Museum on Lake Shore
Drive to attend Saturday morning lectures for
children on natural history. Mrs. Hoogstraal then
convinced museum administrators to allow Harry
to listen also to Saturday afternoon lectures for
1844
H Hoogstraal, Harry
adults. Here they were exposed to fascinating
adventures by speakers such as Admiral Byrd and
makers of wildlife films, and Harry was allowed
into the research area of the museum. This expo-
sure kindled a passion for natural history that
Harry maintained for the rest of his life.
He went on to receive the B.A. degree from
the University of Illinois and, while working on a
M.S. degree from the same university, conducted
four expeditions (1938–41) exploring the distri-
bution of animals and plants (first on reptiles and
amphibians) and later with the late Robert Traub
and Kenneth L. Knight (fellow student) collecting
invertebrates, mammals and birds. During the sec-
ond year, the brakes failed on a truck that Harry
was driving on June 20, 1940, while on a remote
mountain road in the Sierra Madre mountains, in
the central highlands of Mexico. Harry was caught
beneath the truck’s frame when it rolled, while his
associates in the back were thrown clear. His back
was broken and he suffered other injuries and was
taken to the hospital in Mexico City where physi-
cians predicted he would never walk again. How-
ever, two physicians from the University of Chicago
were visiting and they just happened to have a new
sulfa drug, sulfonilimide, that had been recently
developed. He was treated with this drug, making
him one of the first persons to receive it.
After receiving the M.S. degree in 1942 he
joined the U.S. Army and worked on mosquito
­taxonomy in New Guinea and in the Philippines.
During wartime he developed plans for a Philippine
expedition with Donald Heyneman, Floyd Werner
and Philippine colleagues by convincing the herpe-
tologist, Karl Schmidt, Curator of Zoology, Field
Museum to join with the Philippine National
Museum in conducting a faunal survey.
Later, in 1948–49, he conducted surveys in
East Africa and Madagascar under the U.S. Navy
and the University of California African expedition
and became head of the Sudan sub-unit of Naval
Medical Research Unit #3, where he conducted
host-parasite studies in the equatorial province of
the Sudan. This resulted in his 1,099 page mono-
graph on the “Ticks of the Sudan” (1956), which
continues to be cited today. In 1950 he became
Head of the Medical Zoology Department at
NAMRU #3 in Cairo, a post that he held until his
death. He received the Ph.D. from the London
School of Hygiene and Tropical Medicine in 1959,
with the D.Sc. from the same institution in 1971.
He also received a doctorate (honoris causa) in
1978 from Ain Shams University, Cairo and an
honorary D.Sc. from the University of Khartoum
in 1983.
Starting in the late 1940s, Harry pursued the
study of ticks with a single-minded dedication,
­producing over 500 research papers, an 8-volume
bibliography of ticks and tick-borne diseases, and
over 1,750 translations of Russian and Chinese arti-
cles. Harry was interested in ticks on a world-wide
basis, not just in Africa, and few scientists outside of
­Russia or China could read these languages. When
Madam Pospelova-Shtrom published her paper on
a new Argas tick from a vulture in Turkmenia (for-
mer USSR) (previously known as the fowl tick,
Argas persicus), Harry made the translation (T89)
available to anyone working on ticks. He would
routinely send his publications to anyone interested
in ticks, as well as copies of his bibliography, and he
cooperated with Rocky Mountain Laboratory sci-
entists in Montana (USA) as well as tick authorities
in England, and in South Africa. He and Makram
Kaiser then reviewed the Argas persicus problem by
re-examining as many ticks as possible from muse-
ums and collections that had previously been mis-
identified as persicus, resulting in several new
species being described such as Argas arboreus from
heron and cattle egret rookeries in Cairo, Argas
­africolumbae from swallows in South and East
Africa, A. zumpti from a vulture in South Africa,
and A. walkerae from domestic poultry in southern
Africa. In the Americas, persicus were also re-
examined as it was originally believed this tick
occurred world-wide between 40° N and 40° S lati-
tude. Glen Kohls and C. Clifford then reported A.
sanchezi in southwestern USA and Mexico, A. radia-
tus in central and southern USA, and A. miniatus in
Panama and South America; all had previously
been misidentified as persicus.

Harry was considered a workaholic and slept
only short periods at night. However, this rest-
lessness was forced upon him due to the truck
accident in Mexico that left him in considerable
pain. Another reason for Harry’s tremendous
productivity was that he took full advantage of
postal services (before e-mail) to persuade any-
one working in tropical countries to send him
ticks they had collected. He also kept up-to-date
with his correspondence (150–200 letters/week),
which often kept three or four secretaries work-
ing full time to keep up with him. Harry would
have the ticks sent to him identified and cata-
logued, or refer them to Kaiser in Cairo or Clif-
ford and Keirans (in Montana), and then
correspond with the collector regarding their
identification, often requesting more material.
For example, Dr. Alex Fain (Belgium) sent Harry
males and larva of an argasid tick from fruit bats
that were collected in Zaire (1956) but Harry
needed the other tick stages so when the late Dr.
Robert Usinger, on sabbatical leave from Univer-
sity of California was planning a trip to Africa to
collect cimicid ­Heteroptera, Harry asked him to
look for more material of this tick; subsequently,
Usinger collected females and nymphs that
turned out to be a new species, Ornithodoros
(now Carios) faini, from roosts of Rousettus fruit
bats in the Congo. Usinger sent a sample of the
live ticks to Harry in pouch-mail from Zaire and
hand carried another sample to Cairo to ensure
their survival for rearing purposes so that all of
the stages (larva, nymph, male, female) could be
described and illustrated in the published manu-
script. Harry, as well as his colleagues in Mon-
tana, didn’t wish to describe new species of ticks
based on only one of the stages. When a particu-
lar tick was collected it would be catalogued and
compared with closely related material that was
often laboratory reared. Previously, tick workers
had described ticks based on finding one sex (the
male, for example) and confusion could later
result when other stages were collected. Harry
would not describe a species based on only a sin-
gle specimen.
Hoogstraal, Harry
H 1845
Harry also did not believe in what he called
“tampering with present concepts of tick genera,”
something done in present day with the discovery
of RNA/DNA and molecular studies. He believed
that phylogenetic trends among ticks, especially in
the genus Argas, were lucidly demonstrated on a
subgeneric level.
A portion of the ticks collected would always
be preserved and portion “ground up” by his asso-
ciates in Cairo or at the Rocky Mountain Labora-
tory (Montana, USA) and tested for arbo-viruses.
This was undertaken before the tick arbo-viral
activities of the U.S. Public Health Service were
phased out.
Robin Rice (Hawaii) was an individual Harry
recruited to collect ticks for him on Peruvian cliffs,
in the Galapagos of South America, in coastal
Texas of the USA, and in southeastern Kenya of
East Africa. Some of the African material that Rice
collected remains undescribed to this day, as more
material was needed by Harry before publication
to be sure that they were new species. This author
(E.R.E) accompanied Harry to an escarpment
south of Nairobi near Lake Kwenia to search for
argasid ticks of the Griffon vulture (1975). There
we examined the massive thorn-stick nests of the
red-billed buffalo weaver in Baobab trees (Kiboko
Range Research Station) for a tick previously col-
lected there by Rice and tentatively known as
A. bubalornis; it remains undescribed.
His enormous tick collection was combined
with the Rocky Mountain Laboratory collection
and his material is currently housed at the U.S.
National Tick Collection, currently on loan with
Georgia Southern University in Statesboro.
Harry was a complex man, having many inter-
ests, including sculpture and culture of cacti. He
died in his sleep in Cairo on his 69th birthday, on
February 24, 1986.
References
Heyneman D (1985) Presidential address. Introduction of
President Hoogstraal. In: 60th Annual Meeting of the
1846
H Hooktip Moths (Lepidoptera: Drepanidae)
American Society of Parasitologists, Athens, Georgia,
pp 685–686
Keirans JE (1986) Harry Hoogstraal (1917–1986). J Med
Entomol 23:342–343
Linsley EG, Gressitt JL (1972) Robert Leslie Usinger: autobi-
ography of an entomologist. In: Memoirs of Pacific
Coast Entomological Society, San Francisco, CA,
pp 223–224
Yunker CE, Keirans JE (1998) Makram N. Kaiser 1930–1996.
Exp Appl Acarol 22:61–62
Hooktip Moths (Lepidoptera:
Drepanidae)
john b. heppner
Florida State Collection of Arthropods,
Gainesville, FL, USA
Hooktip moths, family Drepanidae, comprise 812
species worldwide, but predominately Oriental
(647 sp.); none are known for the Neotropics and
only a few are in the Nearctic; the actual fauna
probably exceeds 950 species. Three subfamilies
are known: Drepaninae, Oretinae, and Nidarinae
(the latter only from Madagascar). The family is
in  the superfamily Drepanoidea, in the section
Cossina, subsection Bombycina, of the division
Ditrysia. Adults small to medium size (18–66 mm
wingspan), with head scaling normal; haustellum
small or absent in some species; maxillary palpi
Hooktip Moths (Lepidoptera: D ­ repanidae),
­Figure 40  Example of hooktip moths
­(Drepanidae), ­Tridrepana flava (Moore) from
Taiwan.
vestigial; antennae bipectinate to filiform; body
sometimes somewhat robust. Wings broadly tri-
angular, usually with the forewings falcate; hind-
wings mostly rounded (Fig.  40). Maculation
mostly shades of brown to yellow with various
markings, sometimes more colorful, or hyaline to
white with tan bands. Adults nocturnal. Larvae
are leaf feeders. Host plants include a variety of
plants in Anacardiaceae, Betulaceae, Caprifoliaceae,
Fagaceae, Myrtaceae, Rosaceae, Theaceae, and others.
A few are economic.
References
Bryner R (1997) Drepanidae – Sichelflügler. In: Schmetter-
linge und ihre Lebensräume: Arten-Gefährdung-Schutz.
Schweiz und angrenzenden Gebiete, 2:447–476, pl 11.
Pro Natura-Schweizerische Bund fuer Naturschutz,
Basel
Holloway JD (1998) Family Drepanidae. In: The moths of
Borneo, Malayan Nature Society (Malayan Nature Jour-
nal, 52), Kuala Lumpur, Malaysia, 8:17–70, pl 1–4, 6, 8
Seitz A (ed.) 1911–33. Familie: Drepanidae. In: Die Gross-
Schmetterlinge der Erde, 2:195–206, pl 22–23, 30, 48
(1911); 2 (suppl):167–170, pl 10 (1932–33) 6:631–633,
pl 86 (1928); 10:443–490, pl 48–51 (1922–23); 14:287–
292, pl 41 (1927). A Kernen, Stuttgart
Watson A (1965) A revision of the Ethiopian Drepanidae
(Lepidoptera). Bull Brit Mus (Nat Hist), Entomol 1–178
(Suppl 3): 18 pl
Watson A (1968) The taxonomy of the Drepanidae repre-
sented in China, with an account of their world distri-
bution (Lepidoptera: Drepanidae). Bull Brit Mus (Nat
Hist), Entomol 1–151 (Suppl 12):, 14 pl
Wilkinson C (1972) The Drepanidae of Nepal (Lepidoptera).
Khumbu Himal 4:157–332
Hope, Frederick William
Frederick Hope was born in London on January 3,
1797. In 1817 he entered Oxford University, and
graduated in 1820 with a B.A. degree in classics,
and in 1823 obtained an M.A. The intent was that
he should enter the church, and he was presented
with a curacy in the county of Shropshire, but his
health did not allow this to continue. While he was
an undergraduate, he became fascinated with the

natural sciences and spent much time studying
geology and entomology. After leaving Shropshire,
he returned to London and,ww later aided by Ellen
Meredith, the wife he married in 1835, began to
collect natural history objects into his own private
museum. He built his collections not just by his
own collecting, but by purchase, gifts and exchange,
such that he acquired materials from Britain and
abroad. He travelled in France, Germany, Holland
and ­Switzerland in the 1830s. He was elected ­fellow
of The Royal Society in 1834. He was one of the
founder members of the Entomological Society
of  London in 1833, and served as president in
1835–1836, 1839–1840, and 1845–1846. He also
published mainly taxonomic works on insects,
especially Coleoptera, and many in the pages
of  the journals of the Entomological Society of
­London. These works included descriptions of
some of the insects collected by Charles Darwin.
In 1855, Oxford University began to build a new
museum to house scientific collections, and Fred-
erick Hope’s collections and personal library were
promised to that museum by deed of gift, with the
stipulation that a conservator of his collections
should be appointed by the university. Thus, the
position of Hope Professor of Zoology was cre-
ated, and this position was held first by J. O. West-
wood. Frederick Hope died on April 15, 1862. A
few entomologists (faculty, technicians, and grad-
uate students) associated with the Hope Professor
of Zoology (Entomology) remained housed with
the Hope Entomological Collections and Hope
Library in the University Museum, as the Hope
Department of Entomology, until 1978. Then, the
name Hope Department of Entomology disap-
peared, merged into Oxford University’s Zoology
Department. The title of Hope Professor of Zool-
ogy (Entomology) disappeared in 1995. But, the
name “Hope Entomological Collections” still per-
sists for the insect collections housed in the Uni-
versity Museum, and a curator and assistant
curator are responsible for them. The lecture room
of the University Museum, on Saturday June 30,
1860, was scheduled to be the site of the now-fa-
mous debate between Thomas Henry Huxley, a
Hopkins, Andrew Delmar
H 1847
staunch supporter of Darwin, and ­Samuel Wil-
berforce, Bishop of Oxford and creationist.
­Wilberforce “begged to know, was it through his
[Huxley’s] grandfather or grandmother that he
claimed descent from a monkey.” Huxley replied
that he would not be ashamed to have a monkey
as an ancestor, but he would be “ashamed to be
connected to a man who used great gifts to obscure
the truth.” It would be expected that Hope and
Westwood would have been present at that meet-
ing of the British Association. Their reactions were
not recorded. Neither had accepted Darwin’s the-
ory, although both had been correspondents of
his, and both described species that he collected
during the voyage of the Beagle.
References
Irvine W (1959) Apes, angels, and Victorians. Meridian,
Oxford, OH, 399 pp
Smith AZ (1986) A history of the Hope entomological col-
lections in the University museum, Oxford, with lists
of archives and collections. Clarendon Press, Oxford,
xiii 1/2 172 pp
Hopkins, Andrew Delmar
kenneth w. mccravy1, c. wayne berisford2
1
Western Illinois University, Macomb, IL, USA
2
The University of Georgia, Athens, GA, USA
A. D. Hopkins, a pioneering entomologist, is gen-
erally considered the “father of forest entomol-
ogy in North America.” He was born in Harrison
County, Virginia (now Jackson County, West
­Virginia) on August 20, 1857. Though he never
received much formal education, Hopkins
showed an interest in natural history at an early
age, spending many hours collecting and study-
ing rocks, plants, and animal life in the rural West
Virginia landscape where he grew up. His formal
education ended at age 17, though he would
receive an honorary doctoral degree in 1893. As a
matter of fact, Hopkins appears to have had a
1848
H Hopkins, Andrew Delmar

relatively low opinion of formal education, and

espoused the opinion that “college is a place
where pebbles are polished and diamonds
dimmed.” His lack of formal training certainly
did not prevent him from becoming an eminent
scientist and ushering in the discipline of forest
entomology in the United States.
Hopkins was deeply involved in the agricultural
activities of West Virginia, and organized one of the
first farmer’s cooperatives in the state, the Farmers’
Institute Society. He was an accomplished agricultur-
alist himself, developing a superior strain of timothy
hay through plant breeding, and introducing pure-
bred sheep and cattle to the area. Through his activi-
ties with the Society, he became acquainted with Dr.
John Myers, the first director of the West Virginia
Agricultural Experiment Station. In 1889 Hopkins
wrote Dr. Myers, volunteering to be the entomologist
for the Station, but was politely rebuffed due to his
lack of formal training. Not one to be easily discour-
aged, Hopkins wrote back, offering to work on a trial
Hopkins, Andrew Delmar, Figure 41  Andrew
basis for $1.00 a day. Dr. Myers offered him the job on
D. Hopkins, the Father of American Forest
trial for 3 months at $50.00 per month. Hopkins offi-
Entomology.
cially began work on March 1, 1890.
Hopkins early research projects (Fig.  41)
involved a large bark beetle outbreak in the pine an early forest entomology importation biological
and spruce forests of eastern West Virginia, and control program, importing the clerid beetle
Hessian fly as a pest of wheat. His seminal work on Thanasimus formicarius from Germany for release
the Hessian fly showed that delayed planting could against the beetles. However, there was no evidence
control the fly, and he produced appropriate plant- that the clerids became established, and the bark
ing dates for West Virginia and much of the United beetle epidemic subsided following a severe winter
States as well. This work was the foundation for a in 1893–94. Interestingly, Hopkins and his clerid
lifelong interest in bioclimatics, the study of the release received credit for ending the bark beetle
relationships among life, climate, and the seasons. epidemic, even though Hopkins emphatically denied
Hopkins’ discovery of a massive bark beetle that the clerids had become established.
outbreak in the pine and spruce forests of Randolph During his 12 years with the Experiment
County and his subsequent investigations of ­Station, Hopkins published scientific papers and
this group of insects launched his career as a forest bulletins at a prodigious rate. In addition to his
entomologist, and marked the beginning of forest papers on the southern pine beetle outbreak, his
entomology as a distinct discipline in the U.S. It was most important forest entomology works included
for this work that Hopkins would receive his honorary the “Catalog of West Virginia Scolytidae (Bark
Ph.D. degree in 1893 from West Virginia University. ­Beetles) and Their Enemies” and “Catalog of West
After extensive sampling, Hopkins concluded that Virginia Forest and Shade Tree Insects.” As an indi-
the southern pine beetle, Dendroctonus frontalis, was cator of Hopkins’ productivity, during a single year
the major cause of tree mortality. Hopkins initiated with the Experiment Station he described 196 new

species and six genera of insects, and this was pri-
marily background work for his studies of insect
biology and pest control. In addition to his research
work, Hopkins duties included answering hun-
dreds of extension-type questions from citizens on
a wide variety of entomological subjects.
Hopkins expertise in forest entomology was
recognized by the U.S. Department of Agriculture,
and Gifford Pinchot, head of the Division of
­Forestry, hired him to conduct special investiga-
tions of insect problems in forests of the western
U.S. Hopkins had traveled through parts of the far
west in 1899, and made observations on stand
characteristics of ponderosa pines that became the
basis for a risk rating system for this species. In
barely over 2 months, Hopkins correctly identified
and detailed most of the important forest insect
­problems of the far west. In October 1901, he
was sent to the Black Hills region of South
Dakota to investigate bark beetle problems there.
Though he was in the Black Hills for only 4 days,
Hopkins ­collected an incredible amount of infor-
mation on the mountain pine beetle, Dendroctonus
ponderosae, which he published in a bulletin only
3 months later.
In 1902, Hopkins took a position with the
U.S. Department of Agriculture Bureau of Ento-
mology in Washington, D. C., initially working
under the famous entomologist C. V. Riley. Within
2 years, however, he was named head of the
­Division of Forest Insects, a position he would
hold for 21 years until his retirement in 1923.
Among those who worked under his direction
during this period were F. C. Craighead, H. E.
Burke, J. M. Miller, F. P. Keen, and W. F. Fiske, a
­virtual “Who’s Who” of forest entomologists in
the early twentieth century.
Characteristically, Hopkins remained quite
active after his retirement from the Bureau of
­Entomology. He quickly resumed pursuit of his
lifelong interests in plant breeding and bioclimat-
ics. One well-known product of this research is
Hopkins’ Bioclimatic Law, which states that
­elevational decrease in temperature is mirrored in
latitudinal decrease in temperature as one moves
H-Organ
H 1849
away from the equator. His Wood County, West
Virginia farm was designated a special field station
of the Department of Agriculture, “The Kanawha
Farms Intercontinental Base Station for Bioclimatic
Research.” Hopkins was married in 1880 to Adelia
Butcher, and they had four children, including W.B.
Hopkins, who became a well-known petroleum
engineer and was listed in “Who’s Who in Ameri-
can Men of Science.” A. D. Hopkins died in 1948,
truly one of the outstanding scientists of his time.
References
Berisford CW (1991) Andrew Delmar Hopkins – a West
­Virginia pioneer in entomology, vol 14. West Virginia
University Agricultural and Forestry Experiment
­Station, Morgantown, West Virginia, pp 20–26
Furniss MM (1997) American forest entomology comes on
stage: bark beetle depredations in the Black Hills forest
reserve, 1897–1907. Am Entomol 43:40–47
Johnson MM (1930) A half century with a West Virginia
scientist. West Virginia Rev November:47, 65
Hopperburn
A plant disease caused by toxicogenic secretions
associated with feeding, usually by leafhoppers
(Hemiptera: Cicadellidae), hence the common
name.
Hoplopleuridae
A family of sucking lice (order Phthiraptera).
 Chewing and sucking lice
H-Organ
serap mutun
Abant Izzet Baysal University, Bolu, Turkey
H-organ is a neurohaemal organ associated with
the perisympathetic organs of the ventral nerve
cord. The H-organ was first discovered in 1986 in
1850
H H-Organ

various Lepidopteran species and later was studied

in some Orthopterans. It is named the H-organ due
to its shape, resembling the letter H. It is located in
the prothoracic region between the subesophageal
ganglion and prothoracic ganglion. It is approxi-
mately three times smaller than a single prothoracic
gland. It extends across the narrow cervical region
and is exposed to good hemolymph flow. The organ
consists of three parts: anterior arms, posterior arms
and a central region. The anterior arms stretch out
laterally toward the ­prothoracic glands and consist
of two types of nerve fibers. The posterior arms are
composed of nerve axons extending backward and
to the lateral arms. The central part of the organ is
connected with the prothoracic ganglion via median
nerves. The organ includes abundant nerve fibers,
invaginated neurilemma, neurosecretory products
and irregularly distributed glial cells. The H-organ
is very similar to the regular metameric perisympa-
thetic organs. However, the organ has a very complex
structure, differing from other regular perisympa-
thetic organs in its appearance, and consisting of
two fused perisympathetic organs. The first belongs
to the subesophageal ganglion attached from its
rear, and the second is in the frontal part of the H-Organ, Figure 42  The H-organ in the
prothoracic ganglion. The most complicated part ­prothoracic region of an insect: Prg = prothoracic
appears in the middle region. In this branching area gland; Ha = anterior arm of the H-organ; Ho = 
between the anterior arms, transverse nerve of the H-organ; Hp = posterior arm of the H-organ;
H-organ (Fig. 42) and the posterior arms, an organ Prtg = prothoracic ganglion; Sog = subesophageal
that seems like the corpus cardiacum was found. ganglion.
The H-organ receives neurosecretory product from
both subesophageal and prothoracic ganglia. and neurosecretory axons to the prothoracic glands.
The H-organ often shows species-specific vari- Studies in some Orthopteran species show that both
ations in its shape. Within the order Lepidoptera, in Locusta migratoria and Acrida bicolor have a trian-
Pieris brassicae and some other members of the gular shaped H-organ composed of a central body
family Pieridae and Nymphalidae, the organ is with its extensions connected to the subesophageal
almost rectangular or square. In Actias selene and ganglion by two thick bundles of nerve fibers, and
several other members of the family Saturniidae, it by a single thick connection to the prothoracic gan-
shows a tracheolized structure projecting from the glion. The organ is connected to the prothoracic
prothoracic ganglion where posterior arms seem glands at the laterals by its anterior arms.
absent. In Barathra brassicae and Noctua pronuba Together with other perisympathetic organs,
(Noctuidae) it has rather narrow anterior and as a translucent and poorly visible structure, the
­posterior arms and a more compact central region. H-organ is important in insect development and it
In this case, the anterior arms are well developed so is the main source of innervation and neurosecre-
that the organ provides neurosecretory products tory material for the prothoracic glands, which

secrete ecdysone, a hormone that stimulates larval
molts and metamorphosis in insects. It has direct
neurosecretory connections with the prothoracic
gland and the prothoracic spiracle, and may
­control metabolism in the prothoracic gland. The
H-organ shows more-or-less a continuous growth
without apparent cyclic changes. This acyclic
growth and other nervous characteristics of the
H-organ suggest that the physiological nature of
the H-organ be predominantly of a nervous rather
than glandular character.
The H-organ is well developed in all instars.
During metamorphosis, the H-organ starts to retreat
gradually, and disappears during adult development.
The middle part of the H-organ appears to be hyper-
trophied during the molting periods, containing
material in the form of clusters or droplets, and then
atrophied during interecdysis. Histofluorescence
studies indicate the presence of catecholamines and
indolealkylamines, and some other neurosecretory
products that are azocarminophilic in their nature.
These are found in the lateral parts of the H-organ
­surrounding the central region, a region called the
corpus prothoracale. The lateral branching area with
the corpus prothoracale is richly supplied with trache-
ase, indicating a high metabolic activity in this region
of the organ. Although the function of the H-organ is
not yet clear, it has been suggested that the central
region of the organ with the corpus p
­ rothoracale is the
most important neurohaemal area among the whole
ventral nerve cord among the studied insect species.
References
Abou-Halawa S, Slama K (1986) A neurohaemal organ in the
prothorax of Lepidoptera. Acta Entomol Bohemoslo-
vaca 83:241–252
Abou-Halawa S (1987) The H-organ and innervation of the
prothoracic glands in Galleria mellonella (Pyralidae,
Lepidoptera). Vestn Cesk Spolegnosti Zool 51:81–84
Abou-Halawa S (1987) Detailed morphological studies on the
H-organ in the larvae of Galleria mellonella (Pyralidae,
Lepidoptera). I. The anatomy of the organ. Vestn Cesk
Spolegnosti Zool 51:161–167
Abou-Halawa S (1987) Detailed morphological studies on the
H-organ in the larvae of Galleria mellonella (Pyralidae,
Hormoligosis
H 1851
Lepidoptera). II. The histological structure of the H-organ.
Vestn Cesk Spolegnosti Zool 51:241–245
Bhargava S, Slama K (1987) Growth of the prothoracic H-organ
in larvae of Galleria. J Insect Physiol 33:55–57
Mutun S, Ober A (1998) The localization and structure of a neuro-
haemal H-organ in Acrida bicolor (Thunberg) and Locusta
migratoria (Linneaus) (Orthoptera). Turk J Zool 22:41–44
Horizontal Gene Transfer
The transfer of genetic information from one
species to another.
Horizontal Transmission
This is movement of a pathogen among members
of a species within the same generation, or within
the same season. In contrast, transmission of a
pathogen between members of an arthropod
species by movement from parent to  progeny
(transovarial or transovum transmission) or
between instars within a generation (trans-stadial
transmission) is called vertical transmission.
Hormoligosis
Subharmful quantities (low, sublethal exposure rates)
of stress agents (e.g., temperature, radiation, minor
injuries, chemicals) can be stimulatory to organisms
in a stressful environment. In entomology, this
usually is seen in the increased development rate of
insects or mites, or increased reproduction, following
exposure to insecticides and reared under subop-
timal conditions. It is derived from the Greek words
“hormo” (to excite) and “oligo” (minute quantities).
Hormoligosis is sometimes used to explain popula-
tion resurgence of insects and mites following
insecticide treatment. It is important to separate
direct physiological (horm oligotic) effects from
indirect effects related to the destruction of natural
enemies by low doses of pesticides; either can cause
increases in abundance. Also, the response of insects
to such stressors may be delayed.
1852
H Hormone

References Europe, Asia and North Africa, and was acciden-

Luckey TD (1968) Insecticide hormoligosis. J Econ Entomol
61:7–12
Cohen E (2006) Pesticide-mediated homeostatic modulation
in arthropods. Pest Biochem Physiol 85:21–27
Hormone
A chemical produced internally by one organ and
conveyed, usually by means of the hemolymph, to
another organ, where it can trigger physiological,
morphological and behavioral changes.
 Adipokinetic and Hypertrehalosemic
Neurohormones
 Ecdysteroids, Endocrine Regulation of Insect
Reproduction
 Juvenile Hormone
 Metamorphosis
 Prothoracicotropic Hormone
Horned Powder-Post Beetles
tally introduced to North America in the 1880s. It
was first reported in Canada in the early 1890s. It
colonized most of the continent, and had reached
Venezuela by the 1930s. Horn fly arrived in Brazil
in the mid 1980s and has spread through much of
the cattle -producing areas of South America.
The adult horn fly lives in close association
with cattle. The fly is about 5 mm long; gray, with
four dark stripes on the top of the thorax. The wings
are broad and held in a V-shape at rest (Fig. 43).
The eyes are large and red, widely spaced in the
females and close together in the males. The
mouthparts are similar to the stable fly’s with a
heavily sclerotized labium that forms the sheath of
the proboscis (Fig. 44). The food canal is formed
by the labrum-epipharynx and the hypopharynx
forms the salivary canal. The labium ends in a pair
of labella that bear sharp teeth used for cutting the
host’s skin. The maxillae are greatly reduced and
only the maxillary palps are visible at the proximal
end of the proboscis. The maxillary palps are lon-
ger than half the length of the proboscis.

Members of the family Bostrichidae (order

Coleoptera).
 Beetles

Hornets
Members of the family Vespidae (order
Hymenoptera).
 Wasps, Ants, Bees and Sawflies

Horn Fly, Haematobia irritans (L.)

(Diptera: Muscidae)

tim lysyk
Agriculture and Agri-Food Canada Lethbridge,
Lethbridge, AB, Canada

The horn fly is a significant pest of cattle through- Horn Fly, Haematobia irritans (L.) (Diptera: Muscidae),
out much of the world. The fly occurs throughout Figure 43  Horn fly, Haematobia irritans.

Horn Fly, Haematobia irritans (L.) (Diptera:
Muscidae), Figure 44  Head and mouthparts of
horn fly.
The adult flies spend most of their lives in
close contact with cattle. Adults are most abundant
on bulls, followed by steers, cows and calves. The
adults are quite mobile and the distribution of flies
on the cattle body will change markedly during
the day as adults move from the backs and sides of
the animals to the belly. The proportion of flies on
the belly shows a distinct seasonal trend, and is
highest in late summer and lowest in the spring
and fall. This movement may be in response to
increasing numbers on an animal, environmental
fluctuations, increasing proportion of gravid
females in the population, and the presence of
lesions along the belly. Both sexes feed on cattle by
cutting the skin with the labellar teeth and ingest-
ing the resulting blood and tissue. Flies can ingest
from 11 to 21 mg of blood per day. The irritation
resulting from feeding activity of a relatively few
flies (12–25 flies per animal) is believed to be
responsible for an immediate decline in animal
productivity associated with behavioral changes
that reduce forage utilization and feed efficiency.
Horn Fly, Haematobia irritans (L.) (Diptera: Muscidae)
H 1853
Horn fly infestations of yearling cattle have been
responsible for 18% reduction in weight gains.
When fly numbers exceed 200 flies per animal,
cattle will undergo physiological changes that
result in further reductions in productivity that
increases with fly numbers. Horn fly infestations
on cows can indirectly reduce weaning weights of
calves by reducing milk production by the cow.
Every 100 flies per cow can reduce calf weaning
weights by 8%, with reported decreases ranging
from 3 to 16%.
Female horn flies use ingested blood for egg
production. Females feed for 2–3 days before ovi-
position, and lay from 8 to 13 eggs per day. Eggs are
laid every 1–2 days until death. As females become
gravid, they move to the rear and underside of the
cattle, and leave to lay eggs on the undersides of
newly deposited cattle manure. Although female
may live a maximum of 21 days, the average female
lifespan is generally <1 week. Females could lay
100–200 eggs if they reach the maximum lifespan,
but few do. On average, a female will contribute
20–25 eggs to the next generation.
The eggs are 1.2 mm in length and reddish-
brown in color. Eggs hatch within a day and the
first instar larvae enter the pat. Larvae feed on bac-
teria within the manure and pass through three
instars. The third-instar larvae are easily identified.
These are typical maggots with a reduced head and
an anteriorly tapering body. The posterior spira-
cles have a characteristic pattern with a button
located on the inside margin surrounded by three
sinuous slits. Larval survival is influenced by
chemical factors such as manure pH, nitrogen
content, and moisture; physical factors such as
temperature; and biotic factors such as predation.
Larvae survive best in feces from animals feeding
on forage compared with those fed concentrated
or grain-based diets, hence the greater occurrence
of this pest in pasture or rangeland systems. The
larval stage is completed in 1–2 weeks depending
on temperature. Larval development requires
9 days at 20°C and 4 days at 30°C.
The pupal stage is formed beneath the pat
(feces). Pupae weight about 6 mg and develop
1854
H Horn Fly, Haematobia irritans (L.) (Diptera: Muscidae)
within a reddish brown puparium formed by the
last larval skin. The duration of the pupal stage is
strongly influenced by temperature, requiring about
12 days at 20°C and 5 days at 30°C. The pupal stage
occupies about 55% of the total immature develop-
mental time. Pupae can be attacked and killed by a
variety of hymenopterous parasitoids. When pupal
development is complete, the adults emerge, find a
host, mate and continue the life cycle.
The entire life cycle is completed in 2–4 weeks
under field conditions. Reproduction is continu-
ous, and there are several overlapping generations.
The number of generations per season increases
with warmer climates. In northern areas, there
may be 3–5 generations per year, and 5–8 genera-
tions per year in southern regions. Populations
generally show a major peak in mid-summer in
northern areas, and may show spring and fall
peaks in southern climates.
In the fall, environmental cues trigger the
production of diapausing pupae, the overwinter-
ing stage. The increasing proportion of diapausing
pupae produced in the fall results in reduced
recruitment to the adult stage and adult popula-
tions decline. Earlier research suggested that dia-
pause is induced by declining photoperiods acting
on female flies causing them to produce eggs that
are predisposed to enter diapause when they reach
the pupal stage, and that temperatures to which
the immatures are exposed mediate the propor-
tion of pupae that enter diapause. More recent
work has indicated that diapause occurs in
response to reduced temperatures experienced by
the larvae during a diapause-sensitive period and
that maternal influences play little role in diapause
induction.
The diapause sensitive period begins when
immatures have completed 10% of their develop-
ment to the adult stage, and ends when 82% of
the immature developmental period is completed.
The incidence of diapause increases as tempera-
tures decline and a greater number of days are
spent in the diapause sensitive period. In warmer
climates, temperatures may not be low enough
for diapause to occur, and generations may occur
the entire year. Temperatures experienced during
the diapause sensitive period are correlated with
overwintering success. Overwintering survival is
low when temperatures average <10°C during
the ­diapause sensitive period, and greater when
temperatures range from 10 to 23°C. Diapause is
not induced at temperatures above 23°C. Once
diapause is induced, overwintering survival is rela­
tively uninfluenced by ambient temperatures. The
diapausing pupae can survive extended periods of
exposure to −5°C, and temperatures below the
pats rarely drop below −10°C, even when air tem-
peratures reach −30°C.
Diapause is terminated gradually throughout
the winter, and post-diapause development begins
in the spring when temperatures increase. The rate
of completion of post-diapause development is
temperature-dependant and adults emerge during
a 3–4 week period in the spring. Flies colonize
cattle, and begin the life cycle again. The flies that
emerge in the spring are produced during a 3–6
week window the previous fall.
Most methods for controlling horn flies are ori-
ented towards treating cattle, as this is the commod-
ity that is affected. The population of adult flies is in
constant association with cattle, and is a convenient
target. Immature stages are typically not targeted for
control as these are protected within the manure pat
and are dispersed throughout the pasture. Chemical
control options are most frequently used and may
be applied to the cattle in several ways. Direct appli-
cations, in the form of sprays or pour-ons can be
used when the animals are gathered. These will pro-
vide 1–2 weeks control of the adults, but after this
period cattle can be re-colonized by newly emerged
adults that were in the immature stages when the
treatment was applied. Reapplication requires gath-
ering the animals for treatment, and this may not
always be practical. Self-application devices can be
placed in the field so that cattle will receive a dose of
insecticide when they contact the application device.
These devices must be placed in areas such as water-
ing sites or salt licks where they cattle will contact
them, and are most effective in forced-use situations.
The most common method for controlling horn

flies is the use of insecticide-impregnated ear tags.
These tags are applied in the spring before cattle are
turned out to pasture, and release insecticides over
the animals’ bodies for the entire fly season. I­ nitially,
these provided season-long control, but horn flies
quickly developed resistance to the commonly used
chemicals. Use of ear tags should be managed to
delay the development of resistance. This can be
accomplished by applying tags to all animals, remov-
ing tags at the end of the season, using tags contain-
ing different classes of chemicals in successive
seasons, and alternating tag use with other applica-
tion methods. Relatively few non-insecticidal
options are available for horn fly control. Various
traps have been designed to remove the adult flies
from cattle when cattle walk through the trap. These
can be effective, especially when used in situations
such as dairies where cattle must walk through the
trap routinely.
References
Lysyk TJ (1991) Use of life history parameters to improve a
rearing method for horn fly, Haematobia irritans irri-
tans (L.) (Diptera: Muscidae) on bovine hosts. Can
Entomol 123:1199–1207
Lysyk TJ (1992) Simulating development of immature horn
flies, Haematobia irritans irritans (L.) (Diptera: Musci-
dae), in Alberta. Can Entomol 124:841–851
Lysyk TJ (1999) Effect of temperature on time to eclosion and
spring emergence of post-diapausing horn flies (Dip-
tera: Muscidae). Environ Entomol 28:387–397
Lysyk TJ, Moon RD (1994) Diapause induction in the horn fly
(Diptera: Muscidae). Can Entomol 126:949–959
Lysyk TJ, Moon RD (2001) Diapause recruitment and sur-
vival of overwintering Haematobia irritans (Diptera:
Muscidae). Environ Entomol 30:1090–1097
McLintock J, Depner KR (1954) A review of the life-history
and habits of the horn fly, Siphona irritans (L.) (Diptera:
Muscidae). Can Entomol 84:20–33
Horn, George Henry
George Horn was born in Philadelphia on April 7,
1840. He was interested in entomology and other
areas of natural history by the time he completed high
Horn, Hermann Wilhelm Walther
H 1855
school. He entered the University of Pennsylvania,
and obtained a medical degree in 1861. In the
American Civil War he served as surgeon to the
infantry of the California Volunteers in 1862–1866.
This allowed him to collect insects in California,
Arizona, and New Mexico. Then he returned to
Philadelphia and excelled in obstetrics. But he was
elected president of the American Entomological
Society in 1866 and, because he was unmarried,
was able to devote all his spare time to the study of
beetles. He obtained greater fame in entomology
than in medicine, publishing 265 papers, and
describing 154 genera and 1,682 species. His ento-
mological papers included syntheses of genera
and higher taxa, and several of them stood for
about a century before being bettered. He was
closely associated with the American Philosophi-
cal Society and was awarded honorary member-
ships in several entomological societies. He died
on November 24, 1897, nearly a year after suffer-
ing a debilitating stroke. His collections are now in
the Museum of Comparative Zoology of Harvard
University.
References
Calvert PP (1898) A biographical notice of George Henry
Horn. Trans Am Entomol Soc 25:i–xxiv
Henshaw S (1989) The entomological writings of George
Henry Horn. Trans Am Entomol Soc 25:xxv–xxii
Herman LH (2001) Horn, George Henry. Bull Am Mus Nat
Hist 265:80–81
Horn, Hermann Wilhelm Walther
Walther Horn was born in Berlin on October 19,
1871. By the age of eight he already was building an
insect collection. In the late summer of 1889 he
made the acquaintance of Gustav Kraatz, who influ-
enced his entomological training. In 1891 he and
his schoolfriend Hans Roeschke published an ento-
mological paper, a monograph of palearctic tiger
beetles, the first for either of them. Following his
father’s wishes, he studied medicine, obtained his
1856
H Horntails
medical degree in 1893, and took his medical prac-
titioner’s examination in 1895. By this time he had
published 39 entomological papers, and in 1896
took his first foreign collecting trip, to Italy, Tunisia,
Algeria, Portugal, and France, lasting 9 months. In
1899, he took his second trip, to Ceylon and India,
lasting 8 months. His third trip came in 1902, to
South, Central, and North America. In 1904, he was
appointed by Gustav Kraatz to the position of direc-
tor of the entomological museum that Kraatz had
founded. In a private house in Berlin, it was first
called Deutsche Entomologische National museum
and in 1920 was renamed Deutsche Entomologis-
che Institut. Walther served in the German army
during World War I, and was posted to the eastern
front. Inflation in Germany after the war caused
great financial difficulties for the institute which,
like some in Britain and the USA, was privately
financed [compare with the American Entomologi-
cal Institute]. In 1922, Walther arranged the
institute’s affiliation with the Kaiser Wilhelm
Gesellschaft zur Förderung der Wissenschaften
[“Kaiser Wilhelm association for the advancement
of sciences”] which made funding more secure, at
least until circumstances changed again. In 1934,
the institute was made part of a national institute
for systematic and applied entomology. Walther’s
taxonomic contributions were almost entirely on
tiger beetles (Carabidae: Cicindelinae), on which he
published 284 papers. His was the work on tiger
beetles in “Genera insectorum” (1908–1915), and in
“Coleopterorum catalogus” (1926). He nevertheless
published 51 papers on insect studies, 19 on museum
studies, and six bibliographies. His (1935–1937)
work “ű ber entomologische Sammlungen, Entomo­
logen und Entomo-Museologie” (on entomological
collections, entomologists, and entomological
museum methods), together with a work coau-
thored with Schenkling (1928–1929) “Index litera-
turae entomologicae” (index to the entomological
literature) were and still are of worldwide impor-
tance to entomo­logists. They are still of worldwide
importance because although computerized bibli-
ographies exist to the literature from some time
(variously) after 1970, the older literature is not thus
computerized, so Horn’s works are  invaluable.
With such interest in bibliographies, he built the
institute’s entomological library to be  the best in
continental Europe. He died on July 10, 1939.
Reference
Korschefsky R (1939) Dr. Walther Horn. Entomol Blätter
35:177–184
Horntails
Members of the family Siricidae (order
Hymenoptera, suborder Symphyta).
 Wasps, Ants, Bees and Sawflies
Hornworm
A caterpillar of the order Lepidoptera, family
­Sphingidae, normally bearing a large, pointed
spine– like structure on the last abdominal
segment.
 Hawk Moths
 Butterflies and Moths
Horse Flies
Members of the family Tabanidae (order Diptera).
 Horse Flies and Deer Flies
 Flies
Horse Flies and Deer Flies
(Diptera: Tabanidae)
james e. cilek
Florida A & M University, Panama City, FL, USA
The family Tabanidae is primarily composed of two
fairly large groups of biting flies known collectively
as horse flies and deer flies. They occur worldwide

and are represented by 4,300 species and subspecies
from 137 genera. Generally, the wings of deer flies
possess a vertical dark band from the mid-line of
the wing down to the margin, while wings of horse
flies can be entirely dark, mottled, or completely
clear. Horse fly wings never possess a vertical band
down the mid-line of the wing. Members of this
family have large eyes; the females are dioptic (eyes
widely separated) and the males are holoptic (eyes
contiguous). Adult size range varies tremendously.
Some deer flies, Chrysops sp., are as small as 6 mm,
while some of the larger horse flies, Tabanus sp., can
be up to 25 mm in length (Fig. 46).
Horse Flies and Deer Flies (Diptera: Tabanidae),
Figure 45  Adult deer fly, showing the vertical
dark band of wing (photo, J. Castner, University of
Florida).
Horse Flies and Deer Flies (Diptera: Tabanidae),
Figure 46  Adult horse fly (photo, J. Castner,
­University of Florida).
Horse Flies and Deer Flies (Diptera: Tabanidae)
H 1857
Only females feed on blood to mature their
eggs, while both sexes feed on nectar sources that
provide carbohydrates. Males cannot bite because
they lack mandibles. Blood-seeking adults can be
severe pests of humans as well as animals.
Tabanids go through complete metamorphosis,
i.e., egg, larva, pupa and adult (Fig.  47). Generally,
the eggs are deposited in layers on vegetation or
other objects hanging over the water (Fig. 48). The
eggs range in size from 1.0 to 2.5 mm and vary in
number from about 100–1,000 depending on the spe-
cies.After oviposition, the female will deposit a water-
proof secretion that covers the eggs to prevent
desiccation and parasitization by wasps. Once the
egg shells harden, they will turn black and some-
times appear as “tar spots” on vegetation. The eggs
hatch in 5–12 days.
Newly hatched larvae will drop immediately
into the water and burrow into the mud to feed
upon organic matter. Developmental habitats for
most horse flies and deer flies are primarily aquatic
to semi-aquatic. However, some species of horse
fly larvae can develop in open pastures or rotted
logs (Fig. 46).
Larva
Egg mass
on leaf Pupa
Adult
Horse Flies and Deer Flies (Diptera: Tabanidae),
Figure 47  Generalized life cycle of Tabanidae.
(Source, redrawn and reproduced from Harwood
and James, and University of Florida).
1858
H Horse Flies and Deer Flies (Diptera: Tabanidae)
Horse Flies and Deer Flies (Diptera: Tabanidae),
Figure 48  Egg mass of Hybomitra lasiophthalma
(source, J. V. Freeman).
The larvae are cylindrical and tapered at
one end. The number of larval instars (stages)
and duration are profoundly influenced by
weather conditions. Several tabanid species
have been reported to be predators of a variety
of invertebrates including insect larvae, crusta-
ceans, snails and earthworms. Sometimes even
small vertebrates can be preyed upon, such as
the unique case of the predatory capture of
adult, newly emerged spadefoot toads, Scaphio-
pus multipicatus, by horse fly larvae, Tabanus
punctifer, on an Arizona mudflat.
In most temperate climates, tabanids usually
have one generation per year. In more southern
regions, a few species may have two generations
per year. But there are exceptions, for some species
take 2 or 3 years to complete development.
Tabanid bites usually cause mild irritation in
most humans. But some deerfly species (Chrysops
sp. and a closely related species Diachlorus fer­
rugatus) have been reported to cause allergic reac-
tions that, in extreme cases, can result in
­hospitalization. Allergic reactions occur as the
body reacts to the anticoagulant injected into the
feeding wound by the female fly in order to stop
blood from clotting and facilitate the uptake of
blood from the host.
Horse flies and deer flies have been incrimi-
nated in the transmission of a number of proto-
zoan, helminthic, bacterial and viral organisms that
cause human and/or animal disease. Some patho-
genic microorganisms, such as the protozoans
­Trypanosoma theileri and Haemoproteus metchnik-
ovi, require the adult fly as a multiplication source
in their life cycle in order to infect animals.
Most disease causing organisms are transmit-
ted by the contaminated mouthparts of the fly.
Several species of horse flies (Tabanus sp.) have
been incriminated in the transmission of Trypano-
soma evansi in Africa, southern Asia and Central
and South America. This protozoan is frequently
fatal to horses, camels and dogs, although it appears
to be non-pathogenic to cattle and buffaloes.
Certain species of deerflies from the genus
Chrysops can transmit the African eye worm, Loa
loa, to humans. Members of the genus Tabanus have
been implicated in transmitting the rickettsial agent
that causes anaplasmosis (Anaplasma marginale) in
cattle. In the southern U.S., a few species of Tabanus
have been found to transmit a viral infection in
horses referred to as equine infectious anemia.
Transmission occurs by the interrupted feeding of
horse flies, via their contaminated mouthparts,
between infected and non-infected horses. Franci-
sella tularensis, the bacterial agent of tularemia, is
sometimes transmitted to man by a few species
of  Chrysops. Tularemia primarily appears during
summer months and occurs in the Northern
­Hemisphere between latitudes 30° and 71°.
Control
Currently, there are no effective long-term meth-
ods to control horse fly and deer fly populations.
Over the years, a variety of non-toxic traps (often
supplemented by a carbon dioxide source as an
attractant) have been developed, but most have
proven suitable only for sampling populations.
Because tabanids are attracted to movement,
it is possible to lure and trap them with suspended
dark spherical objects such as black beach balls

coated with an adhesive. Many tabanids are col-
lected by this method, but it remains to be proven
whether or not they actually reduce local popula-
tions. Box traps are another method (Fig. 49) that
have been used for a number of years to reduce
greenhead adults, T. nigrovittatus, in the salt
marshes around Cape Cod, Massachusetts.
Larval control, through habitat modification
of wetlands or direct application of insecticides to
developmental sites, is not an option because of
state and federal regulatory issues. Insecticide
application against adult horse flies or deer flies
has met with only limited success.
Control of horse flies and deer flies on cattle,
horses and other animals primarily consists of
direct application of over-the-counter insecticides
labeled for such use. However, the effectiveness of
the insecticide to kill tabanids resting on the hair
coat can be reduced by the animal’s grooming
behavior as well as environmental factors, e.g., rain
and/or sunlight. Repellents for humans or animals
Horse Flies and Deer Flies (Diptera: Tabanidae),
Figure 49  Box trap for collecting horse flies on salt
marshes of Cape Cod, Massachusetts (source, J. E.
Cilek).
Horsfall, William R
H 1859
do not offer more than brief relief, at best, from
biting annoyance. Efficacy is often dependent
upon the number of horse flies/deer flies in the
immediate area that are seeking a blood meal.
References
Jackson R, Nowicki S, Aneshansley DJ, Eisner T (1983) Preda-
tory capture of toads by fly larvae. Science 222:515–516
McKeever S, French FE (1997) Fascinating beautiful blood
feeders, deer flies and horse flies, the Tabanidae. Am
Entomol 43:217–226
Mullen GR, Durden LA (eds) (2002) Medical and veterinary
entomology. Academic Press, New York, NY, 720 pp
Horsehair Worms
(Nematomorpha)
A phylum of insect-parasitic invertebrates known
as nematomorphs, Gordian worms, and hair
snakes, the Nematomopha are named after the
tendency of these worms to be long and thin,
resembling the hairs of horses. Also, these inverte-
brates are commonly found in water sources where
livestock such as horses drink. In most cases this is
due to the tendency of common hosts such as
grasshoppers to leap into water troughs and then
be unable to escape. The horsehair worms con-
tained within the bodies of the grasshoppers then
emerge and swim around in the water
 Nematomorphs (Nematomorpha: Gordioida:
Several Families)
Horsfall, William R
William Horsfall was born in the state of Missouri
on January 11, 1908. He received a bachelor’s degree
in 1928 from the University of Arkansas, and a
master’s degree the following year from Kansas
State University. Then he entered Cornell Univer-
sity as a graduate student, and received a Ph.D. in
entomology in 1933. His employment as a teacher
1860
H Horticultural Oil
and researcher began at Cornell, c­ ontinued at the
University of Arkansas, then South Dakota State
University and finally, beginning in 1947, at the
University of Illinois at Urbana-Champaign. But
World War II interrupted his academic work. He
served in the South Pacific as  commander of a
U.S. military malaria survey unit for 3 years. At the
University of Illinois in 1947–1976, he published
five books (including, in 1955, “Mosquitoes. Their
bionomics and relation to disease” and in 1962.
“Medical entomology. Arthropods and human
disease”) and 140 scientific papers, taught medical
entomology and insect bionomics, and won a cita-
tion for outstanding teaching. His research was on
the bionomics of meloids, grasshoppers, chinch
bugs, bean weevils, scarabs, and mosquitoes. He
was an elected fellow of the American Association
for the Advancement of Science. He received
awards of merit from the  American Mosquito
Control Association, the Zoological Society of Fin-
land, the Entomological Society of America, and
the Illinois Mosquito Control Association. After
retirement, he continued to work as a consultant
and to publish. He died in Illinois on November 18,
1998, survived by his wife, Annie Laurie.
Reference
Berenbaum MR (1999) William R. Horsfall. Am Entomol
45:61–62
Horticultural Oil
john l. capinera
University of Florida, Gainesville, FL, USA
Oils have long been used for their insecticidal
properties, including application of oil to plants to
kill plant-feeding species. Most oils used for insect
control are petroleum products, and are also
known as mineral oils. However, oils are also
derived from plants, and usually referred to as
vegetable oils. Historically, the greatest use of oils
was a winter application to twigs and trunks of
fruit trees and ornamental plants, and this product
is called dormant oil. However, more highly
refined, lighter-weight oils have become available
for use during the growing season, and these are
called summer oils. Both mineral and vegetable
oils, properly refined and applied, can be used for
winter or summer applications.
Dormant Oils
The use of dormant oils was popular because it
addressed some difficult insect control problems,
mostly involving aphid and caterpillar eggs, mites
(both adults and eggs), and scale insects (usually
eggs). Also, the treatments could be made during a
relatively slow season (winter) when there was not
much else that could be done to fruit and orna-
mental crops (except perhaps for pruning). Appli-
cation of dormant oil to trees and shrubs before
bud swelling and bud break poses little risk of
injury to the plant, and yet kills many insects and
mites. Both eggs and active forms of small arthro-
pods are susceptible to oil because the oil appar-
ently inhibits air exchange, often by clogging the
air passages (spiracles) through which arthropods
breathe. Dormant oils are relatively heavy-weight
products, and often contain aromatic compounds
such as sulfur that can cause toxicity to vegetation,
hence the timing of application to avoid active
growth by the plant. Heavy oils evaporate slowly,
which is a good characteristic for insect control,
but which can be damaging to plants. Dormant oils
are normally mixed with an emulsifying agent that
allows the oil to mix with water and be dispersed
uniformly. Upon mixing, the oil solution typically
turns white due to actions of the emulsifying agent;
this is the basis for one of the names applied to dor-
mant oil spray, “white oil.” Oil is normally diluted to
about 2% of the spray mixture. Oils are not applied
before winter hardening has occurred, if the
weather is below freezing, or if the plants are begin-
ning to grow. Some plants are sensitive to oil, so the
product label should be consulted for precautions.
 Horticultural Oil
H 1861

Summer Oils such as soybean, canola and cottonseed. Other

sources include neem seed, sesame seed, rose-
Summer oils are lighter in weight, less contaminated mary, and other plants. Like petroleum-based
with residues that may prove phytotoxic, and useful oils, they can cause phytotoxicity. Neem seed is
for many more plants than are dormant oil sprays. well known as a source of azadirachtin, a growth
They are also called superior, supreme or ultra- regulator-based insecticide and feeding deter-
refined oils. Summer oil is often applied to control rent. However, some neem products contain
aphids, whiteflies, leafhoppers, thrips, mealybugs, mostly neem oil and are relatively free of
mites, and the early stages of many other insects, azadirachtin.
usually at concentrations of 1–4%. Damage can
occur when too much oil is applied, the plants
are under water stress (inadequate soil  moisture), Phytotoxicity
the temperature is high (32°C or 90°F), or the
humidity it too high (the oil cannot evaporate). Among the plants that are known to be sensi-
Vegetation with hairy leaves (dense trichomes) may tive to oil are:
be more prone to damage because more oil is held Azalea, Rhododendron spp.
on the leaf. It is advisable to test a few leaves if you Beech, Fagus spp.
are uncertain about the safety of application, and Black walnut, Juglans nigra
wait a few days to see if damage occurs. Summer Douglas-fir, Pseudotsuga spp.
oils are not selective, and will kill most small insects, Hickory, Carya spp.
including beneficial species. However, once they are Japanese holly, Ilex crenata
dry they are no longer toxic, so relative to traditional Juniper, Juniperus
insecticides they are relatively non-toxic to benefi- Cedar, Cedrus spp.
cial insects. Also, they tend to impart a shine or Maple, Acer spp.
sheen to treated plants, so they are popular Redbud, Cercis spp.
because they improve the appearance of ornamen- Smoke tree, Cotinus coggygria
tal species. Spruce, Picea spp.
Also, neem oil is reported to affect several
plants, including:
Stylet Oils Impatiens, Impatiens spp.
Fuchsia, Fuchsia spp.
Lightweight oils also are sometimes recommended Some roses, Rosa spp.
for disruption of plant virus transmission by Olive, Olea europaea
insects such as aphids; such formulations are called Some carnations, Dianthus caryophyllus
stylet oils. The oil interferes with transmission of As noted previously, if there is uncertainty
virus on the stylets of the insects. Such viruses are about phytotoxicity of an oil product to a crop, it
the non-persistent types. Persistent viruses, which is better to test and watch for several days for toxic
are harbored within the insect, are not affected by effects rather than treat an entire tree or crop with
stylet oils. a product of unknown phytotoxicity. If phytotox-
icity is not a problem, then oils are a great pest
control technique to consider for small arthro-
Vegetable Oils pods. Oils are relatively harmless to everything
except small arthropods and some aquatic insects,
Vegetable oils do not really originate with vege- and of course some plants.
tables. The common sources are oil seed crops  Neem
1862
H Horticultural Oil
Horticultural Oil
Petroleum or botanical oil that is used to control
pests on plant.
Host
An organism that provides food and/or shelter
for another organism. In entomology, it also
is  the insect that supports a parasitoid or
pathogen.
Host Feeding
A behavior displayed by a parasitoid when the
adults feed on insects that normally are used for
larval development.
Host Location in Parasitic Wasps
m. l. henneman
University of Bristol, Bristol, United Kingdom
Generally, three steps are recognized in host
location and acceptance by parasitic insects
(parasitoids) which reproduce by oviposition
on or in the bodies of other insects. First, para-
sitoids must locate the proper habitat in which
their host is found. Second, within the habitat
they must find the proper microhabitat. Within
that microhabitat, they must locate the host
itself, and decide whether or not to accept
it  (i.e., whether or not to oviposit), generally
using cues obtained from physical contact
with the host.
Most studies have focused on the last two
stages, orientation toward the microhabitat
and  host location within the microhabitat.
Microhabitat location has been examined using
laboratory equipment such as wind tunnels and
olfactometers – branched tubes in which an
insect is required to make a decision leading to
one of two or more odor sources. A few have
determined specific compounds to which anten-
nal sensilla are sensitive, using a combination of
a gas chromatograph and electroantennogram
(EAG). The chromatograph separates volatile
complexes into constituent compounds, and
the signal as each compound comes into contact
with the insect’s antenna is recorded.
As a result of these studies we can draw sev-
eral  general conclusions about parasitoid odor
­discrimination and learning. First, biochemical
analysis (using chromatography to separate indi-
vidual compounds) shows that feeding by herbivo-
rous insects on their host plants induces systemic
production of specific volatile compounds that
are  not induced by artificial damage alone. Most
commonly found are the two homoterpenes 4,8-
dimethyl-1,3(E),7-nonatriene and 4,8,12-trimethyl-
1,3(E),7(E),11-tridecataene. These have been found
to be produced by several species of herbivore-­
infested plants from agricultural systems, including
lima bean, apple, cucumber, corn, and cotton.
These compounds are induced during feeding
(at least by beet armyworm caterpillars) by a
compound that was isolated from their saliva,
N(17– hydroxylinolenoyl)-l-glutamine, known as
volicitin (Fig. 50).
O
OH
H H
N
O
O
OH H 2N
Host Location in Parasitic Wasps, Figure 50 ­Volicitin,
a compound in the saliva of beet armyworm
­larvae, that induces production of volatiles
­attractive to wasps. (After Alborn, H. T., T. H.
Jones, G. S. Stenhagen, and J. H. Tumlinson. 2000.
­Identification and synthesis of volicitin and related
components from beet armyworm oral secretions.
Journal of Chemical Ecology 26: 203–220.)

Parasitoids, in general, seem to be able to
exploit these specific induced compounds to more
accurately locate their hosts. They orient upwind
preferentially to plants damaged by ­herbivores, over
artificially damaged and undamaged plants. Parasi-
toids also learn odors, which increases their forag-
ing efficiency. They are more responsive and more
accurate in host location when a host is ­associated
with an odor that a ­parasitoid has encountered
before in the ­presence of a host. This is true not only
of plant odors, but is extendible to novel, artificial
odors such as vanilla and chocolate.
Because a reduction in herbivore feeding is ben-
eficial to plants, it has been suggested that systemic
odor production by plants may have evolved to
exploit odor-orientation abilities of the natural ene-
mies of herbivores; in effect, plants are “calling” wasps
to come and attack the insects feeding upon them.
Several authors, however, have also pointed out that
such an effect could also be explained by wound
response. For example, damage accompanied by the
presence of insect saliva could make a plant more
susceptible to infection by a pathogen, and the sys-
temically released compounds are antipathogenic.
Research to this point has not ruled out either of the
explanations, which are not mutually exclusive.
Significantly less is known about the use of
visual cues by foraging female parasitoids. Most par-
asitoids studied are wasps, and wasps are believed to
rely more on odor for orientation, as they usually
have long antennae and small eyes. Research indi-
cates, however, that parasitic wasps orient to, and
learn preferentially, certain colors and patterns asso-
ciated with hosts. For example, wasps may show a
preference for a pattern of feeding damage on a leaf
that is unique to its host. Parasitoids attacking fru-
givorous (fruit-feeding) species may only be attracted
to colors similar to those of the fruits which its host
infests. Within the range of a meter or two, visual
cues are likely more useful than odors, which might
not be as easily traceable in turbulent air currents.
We lack information about the host-locating
abilities of most parasitoids in their natural
­environments, especially at the level of habitat loca-
tion. A few studies that have examined ­microhabitat
Host Plant Selection by Insects
H 1863
location in the field, however, have reinforced find-
ings in the laboratory. But the mechanism by which
a parasitoid flying through a forest locates a small
egg cluster or first-­instar  larva under a leaf, for
example, remains a mystery.
 Tritrophic Interactions
 Arthropod-associated Plant Effectors (AAPEs):
Elicitors and Suppressors of Crop Defense
References
Alborn T, Turlings TCJ, Jones TH, Stenhagen G, Loughrin JH,
Tumlinson JH (1997) An elicitor of plant volatiles from
beet armyworm oral secretion. Science 276:945–949
Dicke M (1994) Local and systemic production of volatile
herbivore-induced terpenoids: their role in plant-carnivore
mutualism. J Plant Physiol 143:465–472
Turlings TCJ, Tumlinson JH, Lewis WJ (1990) Exploitation of
herbivore-induced plant odors by host seeking parasitic
wasps. Science 250:1251–1253
Vet LEM, Dicke M (1992) Ecology of infochemical use by
natural enemies in a tritrophic context. Ann Rev Ento-
mol 37:141–172
Wackers FL, Lewis WJ (1999) A comparison of color-, shape-
and pattern-learning by the hymenopteran parasitoid
Microplitis croceipes. J Comp Physiol A Sensory Neural
Behav Physiol 184:387–393
Host Plant Resistance
Any host plant-related factor that disrupts the host
selection process or causes the plant to be less suit-
able for insect growth, survival or reproduction
than is normal among plants.
 Plant Resistance to Insects
Host Plant Selection by Insects
S. Finch, R. H. Collier
Horticulture Research International Wellesbourne,
Warwick, United Kingdom
Host plant selection by insects is often divided into
“host plant finding” and “host plant acceptance.”
While the two are easy to separate conceptually, in
practice, they are really part of a continuum of three,
1864
H Host Plant Selection by Insects
rather than two, inextricably bonded links. How-
ever, the central link of host plant finding, thought
previously to be governed by volatile chemicals, has,
until now, proved intractable to scientific experi-
mentation. Thus, the focus here is on host plant
selection by insects associated with cruciferous
plants as, since the classical work of Verschaffelt in
1910, most theoretical studies on herbaceous plants
have used the interaction between insects and cru-
ciferous plants as their test system. Such a selection
is logical, as cruciferous (Cruciferae) vegetable and
oilseed crops are of high economic importance and
are now cultivated on large farms in most parts of
the world. In addition, cruciferous plants are ideal
for biological studies, as their chemistry is well
understood and they support pest species from a
wide range of insect orders.
Many researchers have shown that the num-
bers of pest insects found on cruciferous crop
plants are reduced considerably when the back-
ground of the crop is allowed to become weedy,
when the crop is intercropped with another plant
species, or when the crop is undersown with a liv-
ing mulch. Obviously, if placing non-host plants
in the vicinity of host plants reduces the numbers
of insects that actually find their host plants,
then  this could provide a clue as to how insects
find their host plants. It has been suggested that
when the background of crop plants growing in
bare soil is made more diverse by allowing other
non-host plant species to grow in the inter-row
spaces, that the additional diversity “disrupts”
insects from selecting otherwise acceptable host
plants. Such disruption is considered to be medi-
ated through the non-host plants providing (i)
physical obstruction, (ii) visual camouflage, (iii)
masking of host plant odors, (iv) repellent chemi-
cals, or through (v) the non-host plants altering
the physiology of the host plants. Two other sug-
gestions, named by Root (1973) as the “Resource
Concentration Hypothesis” and the “Enemies
Hypothesis,” have also been used to explain why
fewer phytophagous insects are found on host
plants growing in diverse backgrounds than on
similar plants growing in bare soil.
A discussion of the seven hypotheses put for-
ward to date is presented here, followed by a
description of a theory based on “appropriate/
inappropriate” landings, which the authors believe
is the key, or “missing link,” to host plant selection
by phytophagous insects. Finally, the new theory
will be used (i) to discuss the type of information
required to make intercropping, undersowing and
companion planting more successful, (ii) to sug-
gest how insect biotypes could develop, and (iii) to
describe why wild host plants are not decimated
by pest insects.
Description and Discussion of the
Seven Earlier Hypotheses
Physical Obstruction
This hypothesis was used to describe those situa-
tions in which the host plants were, in effect, hid-
den physically by using larger or taller non-host
plants. For example, tall maize plants were used to
protect bean plants from pest infestations. Tall
plants were considered to be effective because they
obstructed the movement of the pest insect within
the cropping system.
It could be argued that there is an element of
physical obstruction when clover disrupts host
plant finding by pest insects of brassica crops as,
to have maximum impact, the foliage of the clo-
ver has to surround much of the host plant.
Although clover growing in such close proximity
to the host plant will obviously obstruct the
searching insects, no suggestions have been made
as to how this mechanism might operate.
However, a mechanism relying solely on
physical obstruction is not supported by recent
findings in which clover plants were desiccated
so  that they retained the same architecture as
the living plants and differed only in color, being
brown rather than green. When the cabbage
root fly (Delia radicum), the diamondback moth
(Plutella xylostella) and the large white butterfly
(Pieris brassicae), were presented with host plants

surrounded by desiccated (brown) clover, the
number of eggs laid did not differ from the num-
bers laid on host plants presented in bare soil.
Hence, the physical presence of the clover was not
sufficient on its own to reduce the numbers of
eggs laid, a reduction occurred only when the
surrounding clover was green.
Visual Camouflage
This hypothesis was based on the two types of
visual stimuli that induce low-flying insects to
land on plants. The first is a directed response to
the color of the plant, which, in most cases, means
green, and the second is an optomotor response
in which landing is provoked by plants “looming
up” along the path of the flying insect. Anything
that competes with such stimuli, such as other
green plants, or raising the height of the overall
background with weeds so that the distance over
which the host plant can be separated from the
background is foreshortened, helps to visually
camouflage the host plants. This makes the host
plants less “apparent” among the foliage of the
non-host plants.
Although many authors showed clearly that
aphids, whiteflies and certain Lepidoptera pre-
ferred plants that stood out against a background
of bare soil, no attempts were made to determine
how such a mechanism might operate.
Masking of Host Plant Odors
The release of “odor-masking” substances into the
air by non-host plant species is considered to con-
fer some protection to the associated host plants.
Although this “associational resistance” seems a
plausible hypothesis, few data have been collected
during the last 25 years to support it.
The possibility that the odor of the host plant
could be masked by that of the non-host plant now
seems much less likely, though not impossible. For
example, host plant selection by the cabbage root
Host Plant Selection by Insects
H 1865
fly was disrupted when its host plants were sur-
rounded by a range of different plants including
the weeds fat-hen (Chenopodium album), fumi-
tory (Fumaria officinalis) and spurrey (Spergula
arvensis); and cultivated plants such as pea (Pisum
sativum), onion (Allium cepa), carrot (Daucus
carota), rye-grass (Lolium perenne), or clover
­(Trifolium). As each of these non-host plants has a
different odor profile, it seems highly improbable
that they would all be capable of preventing an
adapted specialist insect from finding its host
plants. Furthermore, observations made in wind
tunnels revealed that brassica plants growing in
clover were approached by cabbage root flies as
readily as brassica plants growing in bare soil,
indicating that the odors from the clover did not
mask those of the flies’ host plants. More striking,
however, was that the same disruptive effect could
be produced by surrounding the host plants with
plant models made from green cardboard, or sim-
ply with sheets of green paper, neither of which
were releasing plant chemicals. It seems that once
the characteristic host plant chemicals stimulate
the insects to land, the disruption is caused simply
by providing the insects with a greater number of
green surfaces on which to land.
Repellent Chemicals
It is implicit in this hypothesis that the odors given
off by the non-host plants are sufficiently strong to
actually repel the searching insects. It was sug-
gested that the diamondback moth could be
repelled from cabbages by intercropping the cab-
bages with tomatoes (Lycopersicon esculenta) and
that the highly odorous ragweed (Ambrosia
artemisifolia) could be used to repel the cabbage
flea beetle (Phyllotreta cruciferae) from crops of
collard (Brassica oleracea var. acephala). Such sug-
gestions were made to describe why pest insect
numbers were different in the two situations. They
were not based on scientific experimentation.
Whether or not true deterrency is a mechanism
still needs to be proven. Deterrency usually
1866
H Host Plant Selection by Insects
involves highly aromatic plants that often have to
be crushed and tested in small, confined spaces in
the laboratory to show that they are actually capa-
ble of repelling pest insects. As such tests are far
from natural, the validity of using such data dur-
ing the synthesis of new behavioral mechanisms is
questionable. In reality, no experimental evidence
has been produced during the last 15 years to sup-
port the hypothesis that plants produced effective
levels of chemical repellents.
Plants chosen for their odorous nature, such
as French marigolds (Tagetes patula), failed to
deter the carrot fly (Psila rosae) when used as
the intercrop in carrots. In addition, oviposition
by the diamondback moth was similar on Brus-
sels sprouts (Brassica oleracea var. gemmifera)
plants intercropped with plants of sage (Salvia
officinalis L.) and thyme (Thymus vulgaris L.),
two plant species selected for their pungent
odors. Extracts of the essential oils of sage and
thyme were shown to reduce oviposition by the
diamondback moth, but the effect resulted from
contact stimuli and not from repellent volatile
stimuli. Doubtless, many contact chemicals play
a major role during host plant acceptance, but as
these come into play only after an insect has
landed, they are included only in the second
part of this review, which is concerned with host
plant acceptance.
Altering the Profiles of the Host
Plant Odors
While this seems a novel mechanism, it relies upon
the host plants’ inability to metabolize certain
chemicals they take up from the soil, so that such
chemicals, in effect, change the subsequent physi-
ology of the plant. Many claims are made that
African marigolds (Tagetes spp.) planted between
rows of crop plants reduce pest numbers. It is clear
from the earlier discussions that this is unlikely to
be a direct effect of the odors of the African mari-
golds repelling the colonizing insects. However, it
is well known that species of African marigolds
release large amounts of root exudates, which can
be taken up by adjacent plants. It is possible, there-
fore, that any host plant growing in an intercrop
could be affected directly by chemicals taken up
through its roots rather than by having its odor
masked.
To test whether the uptake of such chemicals
was responsible for the differences in plant coloni-
zation, several batches of the host plants were left
in their pots throughout the test periods. As the
effects of the clover were still evident, and often
within minutes of starting an experiment, a generic
mechanism based on the non-host plants (here,
clover) causing physiological changes in the host
plants cannot be supported.
The Resource Concentration Hypothesis
The last two hypotheses are the ones quoted most
frequently. Both were derived from one study in
which Root monitored insect distribution during
three field seasons in pure stands of collard plants
and in single rows of collard plants bounded on
each side by diverse meadow vegetation.
The “Resource Concentration Hypothesis”
states that phytophagous insects are more likely
to find and remain on host plants that are grow-
ing in dense or nearly pure stands. Phytophagous
insects that arrive in a clump of host plants, by
whatever means, and find conditions suitable, will
tend to remain in the area. This “arresting effect”
of host patches will depend upon several factors
such as the size and the purity of the plant stand
and the type of host plant required by the phy-
tophagous insect. In many cases, this accumula-
tion of specialist insects on a concentrated
resource (here, cultivated Brassica plants) will be
sufficient to increase the numbers of phytopha-
gous insects in that locality. Again, this hypothesis
describes simply the effect of changes in the purity
of a host plant stand on insect numbers and does
not include any attempt to develop a general the-
ory to describe how phytophagous insects select
their host plants.

The Enemies Hypothesis
Contrary to five of the earlier hypotheses, which
claim that the differences are due to the direct
effects of the diverse backgrounds on the behavior
of the pest insects, this hypothesis, like hypothesis
5 (altering the profiles of the host plant odors),
proposes that the effects are indirect. In essence,
this hypothesis proposes that lower numbers of
phytophagous insects are found in complex
­environments because predators and parasitoids
are more effective in such situations. Thus, out-
breaks of phytophagous insects are checked early
by the higher numbers of enemies that can be sup-
ported by the diverse resources available in com-
plex environments. Unfortunately, Root found that
the effectiveness of the “enemies” did not differ
significantly between collards grown as pure
stands and those grown in single rows among
diverse meadow vegetation. Nevertheless, Root
pursued this “enemies hypothesis” by discussing
sets of data collected mainly in England. His own,
more extensive data, however, indicated clearly
that the diversity of both predators and parasitoids
was higher in the pure stands, probably because
more prey/host species were also present in that
habitat. Consequently, he concluded that factors
other than natural enemies were responsible for
much of the differences in insect numbers recorded
between simple and diverse habitats. Although
Root himself discounted the “enemies hypothesis,”
many subsequent scientists have championed the
cause of predators, often on the flimsiest of evi-
dence, or without collecting the data necessary to
support their claims.
Conclusions From Recent Work
Although authors have indicated that diverse
backgrounds can affect host plant selection in the
seven ways described above, it is hard, from the
results presented in recent publications, to refute
the more simplistic view that one mechanism is
operating against all species. Recent work
Host Plant Selection by Insects
H 1867
­ ublished by Finch and Collier (2000) included
p
experiments on the cabbage moth (Mamestra
brassicae), the diamondback moth, the garden
pebble moth (Evergestis forficalis), the small white
butterfly (Pieris rapae), the large white butterfly,
the cabbage aphid, the cabbage root fly and the
mustard beetle (Phaedon cochleariae). Despite
these eight test species being from four insect
orders, the ability of each of them to find host
plants was affected adversely, though to differing
degrees, when their host plants were surrounded
by clover. It appeared that differences in the initial
rates of colonization were the factor that regulated
the numbers of phytophagous insects found on
host plants growing in bare soil or clover, as differ-
ences between the two situations were often appar-
ent within minutes of starting an experiment.
Description and Discussion of the
New Theory
Unfortunately, no one has yet developed any of the
earlier hypotheses into a robust general theory,
they are still only hypotheses. Hence, we have gen-
erated our own theory from detailed studies of
insect behavior.
Instead of the seven hypotheses described
previously, we believe that a mechanism that we
have described as “appropriate/inappropriate
landings” is the central link in host plant selec-
tion by insects. In the overall system, the new
theory of host plant selection can be divided into
a chain of actions involving just three links. In
the first link (Link 1; Fig. 51), volatile chemicals
emanating from plants indicate to flying, recep-
tive insects (action 1) that they are passing over
suitable host plants. Once the odor of the  host
plant in the air becomes sufficiently concentrated,
it induces the insect to land (2). In this way, the
volatile chemicals bring the insects into the close
vicinity of the host plants. However, during the
last few milliseconds, when the insects are only a
short (often <1 m) distance away from the plant,
instead of maintaining their directed response to
1868
H Host Plant Selection by Insects
Distance?
7 5
6
4
Host Plant Selection by Insects, ­Figure 51 ­Schematic diagram to illustrate how diverse backgrounds,
here represented by clover (Trifolium spp.), influence host plant finding by the cabbage root fly. Numbers
represent insect actions 1–7 (see text).
volatile stimuli, phytophagous insects switch to a
directed response to green objects, which, in most
cases, means to plant leaves. It is logical that
vision takes over at this stage, as most flying ani-
mals use vision to “pin-point” a suitable object on
which to land. Therefore, insects that fly over
plants growing in bare soil will be stimulated to
land on host plants, the only green objects avail-
able to them (3a), as most phytophagous insects
avoid landing on brown surfaces, such as soil.
When host plants are growing in bare soil, most
landings will be what we have classed as “appro-
priate,” and so, the host plants will, in effect, “con-
centrate” the insects. In contrast, insects flying
over host plants surrounded by clover land in
proportion to the relative areas occupied by
leaves of the host and non-host (3b) plants, as
specialist phytophagous insects do not discrimi-
nate between the two when both are green. Hence,
any landings made on the non-host plant (3b) are
classed as “inappropriate.” The amount of time
the insects spend on the leaves of the non-host
plants before taking off again (4) is governed by
whether the insects receive acceptable or antago-
nistic stimuli through their tarsal receptors. Once
the insects are again airborne, if they are stimu-
lated to land after flying only a relatively short
distance, they could land on a host plant (5, 7). In
all situations, however, the plant on which the
1
2
3a
3b
3c
insect first lands, even if it is a “host plant,” may
not stimulate the insect sufficiently to cause it to
remain on the plant, and the overall process will
be repeated. If this represented the complete sys-
tem, then under “no-choice” situations in the
field, it would just be a matter of time before the
numbers of eggs laid on host plants growing in
diverse backgrounds were similar to those laid
on host plants (3c) growing in bare soil. However,
this does not occur, as there is a second phase to
host plant finding.
This second phase can be illustrated (Fig. 52)
most clearly by data collected from a detailed
study of the cabbage root fly. Before accepting a
host plant as a suitable site for oviposition, recep-
tive female cabbage root flies make, on average,
four spiral flights before laying eggs beside the
plant. Hence, the insects stand a much greater
chance of “losing” the host plant in a diverse back-
ground as, on average, they repeat the initial
appropriate/inappropriate landing procedure an
additional three times. Observations under labo-
ratory conditions showed that for every 100
females that landed on a brassica plant surrounded
by bare soil, 36 received sufficient stimulation
from the plant to be induced to lay eggs. In con-
trast, only seven out of 100 females that landed on
host plants surrounded by clover managed to lay
eggs. Fewer flies managed to lay eggs in this
 Host Plant Selection by Insects
H 1869

1 1

2
2 3

3 4

5 5
36 7

Host Plant Selection by Insects, Figure 52  Schematic diagram to illustrate how diverse backgrounds,
here represented by clover (Trifolium spp.), influence host plant acceptance by the cabbage root fly.
Numbers represent the four (mean no.) leaf-to-leaf flights made by the fly to ascertain whether the plant
is a suitable site to lay its eggs.

s­ ituation, because following each short spiral flight,
a proportion of the flies landed on the leaves of
the  surrounding clover plants. This failure to re-
contact a leaf of a host plant after any spiral flight
prevented the females from accumulating,
within the allotted time, sufficient stimulation
from the host plant to be induced to lay eggs.
Hence, the barrier that this fly faces when its host
plants are grown in diverse backgrounds is not
chemical nor mechanical, but behavioral, simply
because during the innate series of spiral flights,
the fly must ­continue to accumulate more positive
host plant stimuli each time it lands.
The amount of stimulation the female picks
up on each landing is crucial, and this is where the
phase of host plant finding (Link 2) becomes truly
integrated with host plant acceptance (Link 3). In
essence, the complete system really involves find-
ing and refinding the host plant. Obviously, the
insect can re-find the host plant quite easily in
bare soil situations, but not as easily when the
plant is growing in a diverse background of other
plants. The schematic representation shown in the
final figure indicates that the female cabbage root
fly may only have to visit two leaves of a highly
stimulating plant [1] compared to six leaves on a
poorly-stimulating plant [3] before finding it an
acceptable site to oviposit. Other insects, however,
may accumulate sufficient stimuli to keep them
searching [4], but not sufficient stimuli to induce
oviposition and so will fly away. A similar outcome
results when insects visit several leaves but do not
manage to accumulate sufficient stimuli in the
allotted time to be induced to stay [5]. Two other
variations occur when the insects land initially on
a stimulating leaf, but subsequently on a non-
stimulating leaf. It does not matter whether this
leaf is from a host [6] or a non-host plant [7], as
anything that interrupts (Fig. 53) the rate of accu-
mulation of positive stimuli causes the insect both
to abort its attempt to lay and to move elsewhere.
In addition, interspecific competition may also
become important. At any stage during the host
plant selection process, many of the new immi-
grants may not remain on otherwise acceptable
plants if those plants are colonized already by cer-
tain, but not all, of the other insect species present
in the pest complex.
The physiological status of the insect, which
depends partly on its age and also on how long it
has been deprived of a suitable oviposition site,
also has to be superimposed upon this already
1870
H Host Plant Selection by Insects

Stimulation from plant leaves

Threshold to :-
High Intermediate Low
Insect
lays

Accumulated level of stimulation

1 Lay
[1] [2] [3]

[4]

2 Stay Insect
files
[5] away
3 Leave
[6]

[7]

1 2 3 4 5 6

No. of landings on plant leaves

Host Plant Selection by Insects, Figure 53  The number of leaf landings a cabbage root fly may have to
make before accepting a plant as a suitable site for oviposition or deciding to fly elsewhere. The numbers
in [] represent seven possible variations in the pattern of insect behavior.

complex system. With time, phytophagous insects Although the other seven test species men-
tend to become less discriminating in their choice tioned earlier have not been studied in detail,
of oviposition sites. The condition of the plant is records of the movements of the small white but-
also extremely important, as some cruciferous terfly in the field showed that it always made con-
plant species are more highly preferred than oth- tact with several leaves, interspersed with short
ers, and during their phase of exponential growth, flights, before laying an egg. Although the original
many individual plants become highly stimulating author concluded that this butterfly needed some
to insects. However, even when the insect and the flight time to get the next egg ready, these short
plant are both in the appropriate physiological flights could also represent a behavioral repertoire
state, it counts for nothing the moment the insect similar to that of the cabbage root fly. It seems
makes a wrong choice and alights on any green likely, therefore, that the relative differences in the
object other than a leaf of a host plant. This, how- effects that diverse backgrounds have on host plant
ever, is tempered by the fact that when the host selection by the test species may simply reflect the
plant is highly stimulating, the insect has to visit numbers of contacts/re-contacts the insect has to
fewer leaves and so has less chance of making an make to accumulate sufficient positive stimuli to
inappropriate landing. In addition, the highly lay eggs. Results from recent experiments indi-
stimulating plants invariably induce the individual cated that the diamondback moth was the species
insects to lay more eggs. Detailed descriptions of affected least by diverse backgrounds. It raises the
the multitude of other factors involved during question of whether the diamondback moth has
host plant acceptance can be found in many of become such a major pest of cruciferous crops
the papers published in 1999 in the proceedings simply because it has a limited behavioral reper-
of the Tenth International Symposium on Insect- toire prior to oviposition, and so, lays eggs on more
Plant Relationships. or less the first host plant leaf it encounters.

General Discussion
The “appropriate/inappropriate landing” theory
(Fig. 53) can be used to explain why certain aspects
of host plant finding by phytophagous insects, sup-
posedly regulated by volatile plant chemicals, proved
intractable to scientific experimentation in the past.
Compared to our simple theory of host plant
finding, other theories invoke complex processes
involving volatile chemicals to guide phytophagous
insects to their host plants. For example, it is known
that the odorous environment of a given insect is a
shifting maze of overlapping active spaces. There-
fore, it was suggested that it is in this shifting maze
that the insect must find the active space containing
those few signals that will lead it to its host plant.
However, in the open air, it is turbulence rather than
diffusion that determines the distribution of odor-
ous molecules, and there is nearly always a prevail-
ing wind to blow the odorous molecules away from
the plant. Registering directional cues from odor-
ous molecules is even more complicated for flying
insects, as the movement of the air relative to the
insect depends on the insect’s own activity, and the
wind direction has to be determined by the insect
in the absence of fixed markers.
The evidence that volatile chemicals are the
main regulatory stimuli in the central link of
host  plant finding is weak, as the maximum
­distance recorded for insect orientation to host
plant volatiles in the field is only a few meters. In
wind tunnel experiments, cabbage root flies
flew upwind to a cruciferous plant odor released
at 2.5 g/day, a rate similar to that dispensed from
field traps. This rate of release is at least 105 times
higher than the amount of chemical released
from a healthy cruciferous plant. Although the
flies moved upwind in response to this odor,
<10% of the flights lasted more than 0.5 m. While
the shortness of such flights was described as
unexpected, it would not have been unexpected
had the chemicals involved been regarded as
arrestants rather than attractants. Similarly, with
insect traps releasing large amounts of chemical
to provide directional cues in the field, many
Host Plant Selection by Insects
H 1871
insects miss the trap and subsequently fail to
enter, suggesting again that stimuli other than
volatile chemicals take over once the insect nears
the source of the odor. The current mechanism
seems much more robust than one based on
­volatile chemicals, as the sooner an insect lands,
the sooner it is freed from the plethora of prob-
lems it faces while still in the air.
The “appropriate/inappropriate landing” the-
ory works equally well for generalist feeders, where
the decision of whether to stay is determined pri-
marily by the chemicals the insect detects via its
contact chemoreceptors once it has landed on a
leaf. However, it should be remembered that if an
insect that is considered to be a “generalist” has
been stimulated to land by volatile chemicals
released from a specific plant species, the insect
may continue its search for such a plant rather than
choosing the first acceptable plant it encounters.
Finally, the “appropriate/inappropriate landing”
theory appears to apply equally well to nocturnal
insects, as oviposition by the diamondback moth
and the cabbage moth was not disrupted when their
host plants were surrounded by brown clover.
Although both of these moth species are considered
to be nocturnal, much of their oviposition activity is
concentrated at, or shortly before, dusk. The above
results indicate that during this period, both species
of moth appeared to be able to discriminate between
green and brown objects.
From a crop protection standpoint, the more
non-host plants removed from any crop area, the
greater chance an insect has of finding a host plant.
Hence, current cultural methods are exacerbating
pest control problems, as “bare soil” cultivation
ensures that crop plants are exposed to the maxi-
mum pest insect attack possible in any given
locality.
Future Work
If the theory based on “appropriate/inappropriate
landing” is accepted, it raises searching questions
regarding several aspects of entomological research.
1872
H Host Plant Selection by Insects
In the first instance, we have expressed con-
siderable doubts about whether host plant volatile
chemicals on their own are capable of guiding
phytophagous insects to their host plants. Most of
the detailed experiments with host plant volatile
chemicals have been done by releasing plant odors
from a point source sited at one end of a wind-
tunnel, introducing responsive insects and then
recording whether the insects move upwind to the
source of the odor. Results from this approach
have been disappointing. Often, the only way to
obtain data is to place a visual stimulus, normally
a green object, alongside the site where the volatile
chemical is being released.
If, as the “appropriate/inappropriate landing”
theory suggests, it is only the number of green
objects surrounding a host plant that reduces
­colonization by pest insects, then it should not be
too difficult in the future to quantify the type and
number of plants needed for the diverse back-
ground to reduce pest insect numbers in any given
crop. If several plant types proved appropriate for
a given cultivated crop, it would then simply be a
case of choosing the one that caused the least
reduction in yield to the harvested product.
There is also a need to obtain a better under-
standing of “companion planting” a practice used
frequently by organic growers. Recent data show
that there is no scientific evidence proving that the
odors from highly aromatic plants can actually
deter pest insects. This, therefore, brings into ques-
tion how these aromatic plants produce their effects.
A survey of the literature should help to show
whether such systems are effective only when the
companion plants possess relatively large amounts
of foliage when compared to the crop plants. If this
idea can be substantiated, the differences recorded
may simply be a reflection of appropriate/inappro-
priate landings, and, again, have little to do with
volatile chemicals, no matter how pungent the plant
odors might appear to the researcher.
Apart from its impact in practical pest control
situations, the “appropriate/inappropriate landing”
theory helps also to explain why wild host plants
growing among other plants in natural vegetation
are rarely decimated by pest insects. However, some
insects that are considered to be pest species do
develop on wild-host plants. The question then, is
do such insects prefer to remain on the wild host
plants in subsequent generations, a situation that
could give rise to biotypes? The answer to this ques-
tion is of considerable practical importance. Future
work is required to determine what proportion of
any given pest population develops on wild-host
plants, and whether such insects readily switch back
to the cultivated crop plants. If they do not, then it
should be possible to control certain pest insects by
isolating new crops from earlier infestations.
Additional work is required also to determine
whether appropriate/inappropriate landings influ-
ence the overall distribution of the parasitoids of
the pest species. It is well documented that such
insects use both host plant chemicals and kai-
romones to locate their host insects. Research in
the 1940s in which cabbage plants infested with the
larvae of the small white butterfly were placed in
bare soil crop fields and in hedgerows, showed
clearly that one specific parasitoid, Apanteles rube-
cula, was also affected adversely when the plants on
which its host insects were feeding were placed
into a diverse background. Further work is needed
on other parasitoids to determine whether this is a
general phenomenon. If it is, then the suggestions
of some researchers that diverse backgrounds have
adverse effects on pest insects and no effect on their
parasitoids warrant further study.
Similarly, with respect to the “Enemies
Hypothesis,” it should not be too difficult to show
whether predation is higher on infested plants
surrounded by non-host plants, than on similarly
infested plants surrounded by bare soil.
As much of the aforementioned work has been
based on detailed information on the cabbage root
fly, work is needed to determine the detailed activ-
ity of each of the other pest species. It appears that
diverse backgrounds may have a greater effect on
the cabbage root fly than on the diamondback
moth simply because, before acquiring sufficient
stimulus to oviposit, the cabbage root fly repeats
the sequence of host plant finding four times, but

the diamondback moth only once. It would be dif-
ficult to prove that once the cabbage root fly starts
one of its spiral flights it is again stimulated to land
(arrested) by volatile chemicals, as the distances
between taking-off and landing again are extremely
short. However, olfaction is considered to be used
by phytophagous caterpillars in choosing their
feeding sites, so the intervention of olfactory stim-
uli again at this stage cannot be ruled out.
The hypothesis of “appropriate/inappropriate
landings” does appear to provide a robust descrip-
tion of host plant selection by insects under a
wide  range of different conditions. It shows how
elements of all the earlier hypotheses can be incor-
porated into the overall system, and in ­particular,
how the lack of detailed research on insect behav-
ior, and in particular, on visual ­stimuli, has led to
many of the current anomalies. Apart from the
future work proposed, it will be interesting to
determine whether the same theory regulates host
plant selection by specialist insects found associ-
ated with plant families other than the Cruciferae.
While we believe the simplicity of our theory
makes it all-embracing, only time will tell whether
our optimism is justified.
References
Altieri MA (1994) Biodiversity and pest management in agro-
ecosystems. Haworth Press, New York, NY, 185 pp
Anon (1999) In Menken SBJ, Minks AK Schoonhoven LM
(eds), Simpson S, Mordue (Luntz) AJ, Hardie J (guest eds),
Proceedings of the tenth international symposium on
insect-plant relationships. Entomol Exp Appl 91:265 pp
Finch S, Collier RH (2000) Host plant selection by insects – a
theory based on “appropriate/inappropriate landings”
by pest insects of cruciferous plants. Entomol Exp Appl
96:91–102
Kennedy JS (1978) The concepts of olfactory “arrestment” and
“attraction”. Physiol Entomol 3:91–98
Root RB (1973) Organization of a plant-arthropod associa-
tion in simple and diverse habitats: the fauna of collards
(Brassica oleracea). Ecol Monograph 43:95–124
Schoonhoven LM, Jermy T, van Loon JJA (1998) Insect-plant
biology – from physiology to evolution. Chapman &
Hall, London, UK, 409 pp
Thorsteinson AJ (1960) Host selection in phytophagous
insects. Ann Rev Entomol 5:193–218
Host Specificity of Weed-Feeding Insects
H 1873
Host Specific
An organism that is monophagous, feeding only
on a certain host.
Host Specificity
The degree of specificity of an animal preying
on  hosts. Levels of host specificity include
monophagous, stenophagous or oligophagous,
and polyphagous.
Host Specificity of Weed-Feeding
Insects
luca fornasari
Montpellier, France
The understanding of evolutionary processes is
necessary when considering biological diversity
and the relationships existing between insects
and plants. The major forces in structuring
the  populations of phytophagous insects act
­vertically through trophic levels (e.g., from
­natural enemies of phytophages to phytophages,
and from these to host plants), rather than
­horizontally through interspecific competition.
Therefore, due to the selection pressure by
­phytophagous insects, plants evolved – and some-
times co-evolved with them – interspecific nega-
tive or positive interactions. These are expressed
by a gradient of interactions. At one end of the
spectrum, negative interactions are seen in
defense mechanisms; at the other end of the
spectrum, positive interactions can be seen in
obligate mutualistic interactions. Both are a
result of reciprocal, repeated intimate adapta-
tions and counteradaptations. This is exempli-
fied in phototoxic plants, which developed very
effective defense mechanisms, and in ento-
mophilous plants, which conversely are extremely
specialized and could not even survive without a
given species of insect.
1874
H Host Specificity of Weed-Feeding Insects
Adaptations of Insects to Plants,
and of Plants to Insects
The adaptations in insect-plant relationships
involve behavioral, phenological, physiological,
morphological and biochemical traits of phy-
tophages and plants, which ultimately determine
the success and fitness of species. The attack by
herbivores induces production of allelochemicals
in plants. In turn, the selection pressure driven by
plant resistance, especially through allelochemi-
cals, results in patterns of specialized insect
­natural enemies, whose success is enhanced by
the reduced competition for food that they
encounter once they overcome plant defenses.
These evolutionary adaptations especially involve
the feeding and oviposition habits of phytopha-
gous insects, leading to a high level of specializa-
tion for their host plants, and also defense
mechanisms that allowed them to be resistant.
Resistance is expressed as being non-palatable,
repellent, digestion-inhibiting, or toxic to other
species of insects. Besides toxic chemicals, plant
defenses also include physical (morphological)
structures, and low levels of available nitrogen, or
water/nitrogen ratios. On the other hand, insects
develop specific morphological, biochemical and
behavioral ­adaptations, and synchronize their life
histories with their hosts.
Host Selection
The ability of specialist weed-feeding insects
to  identify, locate and select the proper host as
breeding substrate, is critical to the performance
and survivability of a given species of insect. All
phytophagous insects show some degree of selec-
tivity for their food-plants, and possess a definite
host range. The host selection behavior includes a
specific sequence of activities and responses to
environmental cues, principally orientation, feed-
ing and/or oviposition. Adults usually must choose
the right host for oviposition, thus facilitating
the development of immature stages, which often
have limited mobility. Nevertheless, also immature
stages of insects are capable of displaying host
selection behavior. For oviposition, adults may not
only choose the host plant species, but also a suit-
able individual plant, the most favorable geo-
graphical area, the site, the habitat, plant density,
and the size and the part of the plant. Even subtle
factors such as the degree of protection from biotic
or abiotic damage to eggs, the correct micro-envi-
ronmental conditions for the development of eggs
and immatures, and the nutritional status of the
food supply affect host selection. The spatial pat-
tern of the host plant can affect oviposition behav-
ior, and plant shape can modify its attractiveness
to insects. Adults also regulate their oviposition
behavior in terms of the number of eggs ovipos-
ited per plant, and in some cases will avoid ovipo-
sition on plants where other eggs have already
been laid. This behavior facilitates a regular distri-
bution of eggs on the plants, and regulates the
effect of insect feeding on the host plants. In some
species, adults are deterred from laying eggs if the
plant has already been fed upon by other insects.
The selection of a suitable species of plant by
phytophagous insects is guided by a combination
of physical and chemical stimuli. The sensorial
selection of the host plant by insects involves the
senses of smell, sight, touch, taste and other forms
of contact chemoreception, and may be facilitated
by activity and behavioral patterns typical for a
given species of insect (e.g., pattern of flight).
Plant-produced volatile semiochemicals such as
kairomones and allomones, as well as nonvolatile
secondary chemicals and interactions with micro-
organisms, play a major role in determining insect
behavior. The sense of smell is highly developed in
insects, and the olfactory signal is usually the main
indicator of an appropriate host, often comple-
mented by visual elements. Leaf surface contact
cues are then important in the host selection pro-
cess. Insects also respond to the hardness of tissue,
which is usually associated with lessened feeding.
In general, the balance between phagostimulatory
and deterrent compounds contained in the
ingested plant tissues determines the acceptance

or rejection of the plant by polyphagous insects. In
contrast, for monophagous and many oligopha-
gous species, the perception of a single, specific,
chemical, or group of chemicals of the plant seems
to trigger feeding. A complex of factors and inter-
actions may ultimately cause the association of the
herbivore with the target plant. The characteristics
of the plant causing it to be unsuitable for insects
are described as “xenobiosis” (also called antibio-
sis); the subset of plant characteristics that cause
an insect not to accept a prospective host plant are
described as “antixenosis” (nonpreference).
Learning
Experience may play a role also in the host selec-
tion process, although differences seem to exist
between polyphagous and oligophagous insects. It
is necessary to consider the importance of the
physiological conditions of plants and insects,
­abiotic factors, presence of other plants species and
other biotic factors, as these can greatly influence
the behavior of the insect during the host selection
process, and largely modify the host range that the
insect has under standard conditions.
Plant Defenses
The great genetic plasticity and biological diversity
in insects is amply seen in their specialization on
the most diverse species of host plants. This is clearly
shown also by insects that attack weeds. The species
of weed-feeding insects have a wide array of rela-
tionships with their food-hosts and several mecha-
nisms to overcome plant resistance to attack. Most
phytophagous insects are relatively specific for their
hosts, and generalists represent a minority. How-
ever, it is possible to distinguish different levels of
specificity, in a continuum of patterns of resource
use, ranging from species that are strictly monopha-
gous, to polyphagous species. The degree of speci-
ficity varies, with exceptions, among orders of
insects, but also among lower taxonomical levels,
Host Specificity of Weed-Feeding Insects
H 1875
including species and even sub-specific entities. The
host range depends on characteristics intrinsic and
extrinsic to a species of insect. In the first category
fall all the biological, biochemical and ecological
characteristics of the weed-feeding insect and its
responses to environmental stimuli, whilst in the
second category fall the characteristics related to
the life history, phenology and physiology of the
species of plant considered. For instance, because of
the life history of the plant considered, a species of
insect may not be present at the time of the year
when the plant, or plant part, is susceptible of being
attacked. The fact that narrow host specificity is
prevalent among phytophagous insects means that
most of them developed mechanisms allowing to
overcome hurdles to phytophagy in the food web
represented by the defensive evolutionary barriers
of plants. Land and aquatic plants, including weeds,
show a great biochemical diversity, growth forms,
leaf shapes and seasonal phenologies, and the recip-
rocal (coevolutionary) influences between these
diverse forms of plants and insects, led to the speci-
ficity of weed-feeding insects for their hosts. This
was achieved in various ways by specialized insects.
They alternatively developed the ability to (i) detox-
ify and metabolize xenobiotic compounds con-
tained in the tissues of the host plant; (ii) sequester
toxic secondary plant metabolites; (iii) rapidly
excrete the toxicants ingested; or (iv) attack poison-
ous plants without ingesting an overdose of the
toxic allelochemicals, as a result of behavioral
defenses. On the whole, specialists are more effi-
cient in the use of resources and better competi-
tors than generalists. Specialized monophagous
and oligophagous insects, rather than polyphagous
insects, have the major effect in the regulation of
the abundance of their host plants.
Weed-Feeding Insects
Host specific weed-feeding insects can be used as
beneficial organisms in programs for the biologi-
cal control of weeds, as an alternative to chemical
herbicides, or other forms of control. Therefore,
1876
H Host Specificity of Weed-Feeding Insects
they may be a component of integrated pest man-
agement programs and in some cases may repre-
sent the best, if not the only effective solution to
control exotic weeds introduced in a new geo-
graphical area. This is the case in particular for
rangeland weeds, and weeds that represent an
environmental problem, for example in protected
areas and where the area infested is very large.
Nevertheless, biological control can be success-
fully applied also against weeds infesting crops.
This form of control offers many important
advantages, being effective, selective, environ-
mentally sound, self-perpetuating and cost-effec-
tive, especially in the long term. Information
about the host range of a weed-feeding insect is
essential for organisms that have prospective use
as biological control agents. It is necessary that, in
addition to being effective in controlling the tar-
get plant, it does not harm other plants such as
agricultural crops, ornamental plants, or plants of
ecological value. Therefore, studies must be con-
ducted to be able to predict the effectiveness of
the biological control agent, and to determine
which plants are likely to be attacked in the region
where the insect should be released. Several
approaches were considered to study the level of
specialization of a candidate agent in order to
define its host range and in general these methods
include tests on (i) species of plants taxonomi-
cally related to the target plant for biological con-
trol; (ii) plants attacked by other species of insects
related to the candidate agent; (iii) cultivated
plants of economic importance in the region of
release; (iv) threatened, endangered, or other
native plants of ecological value present in the
area of release; (v) plant species with an increas-
ing phylogenetic distance from the target plant;
and (vi) unrelated plants with biochemical, or
morphological characteristics in common with
the target plant. The evaluation of the safety and
future performance of candidate biological agents
in the release environment depends on a complex
of biotic and abiotic factors and has to be consid-
ered for each specific situation. In order to assess
the degree of specificity of a species of insect,
studies can be conducted in the laboratory and in
the field on the test plant species. The ability of the
insect to feed, produce fertile eggs and oviposit on
the test plants is verified. Also, the ability to com-
plete its development under no-choice and multi-
ple-choice conditions is studied. Once this
information is available, studies on the following
generations can show if the test plant affects their
biology, and if there are variations in fertility, or
other key biological traits. ­Studies conducted in
the field will complement the information
obtained in the laboratory and will help to define
the true host range of the insect. Depending on
the nature of the weed problem, it may be neces-
sary to use organisms with various degrees of spe-
cialization for their hosts, ranging from highly
specific (e.g., attacking exclusively a variety of the
weed species), to less specific insects. In-depth
studies on the life history of these insects are also
required, since often these weed-feeding insects
being considered as candidate agents for biologi-
cal control are new to science, and even when they
are not new species, their biology is ­usually very
poorly known. Due to intraspecific variability
among insects, host range studies should include
a sufficiently large sample of insects to be repre-
sentative of that species, and it is preferable that
they are conducted on precise, geographically dis-
tinct populations. The biology, habits, degree of
host specificity and, if necessary, genetics of pop-
ulations of different origin are compared.
 Foreign Exploration for Insect that Feed on
Weeds
 Biological Control of Weeds
 Arthropod-associated Plant Effectors (AAPEs):
Elicitors and Effectors of Crop Defense
References
Ananthakrishnan TN, Raman A (eds) (1988) Dynamics of
insect–plant interactions. Oxford & IBH Publishing Co,
New Delhi, India
Barbosa P, Letorneau DK (eds) (1988) Novel aspects of
insect–plant interactions. Wiley, New York, NY

Dettner K, Bauer G, Völkl W (eds) (1997) Vertical food web
interactions: evolutionary patterns and driving forces.
Springer, Berlin, Germany
Hunter MD, Ohgushi T, Price PW (eds) (1992) Effects of
resource distribution on animal–plant interactions.
Academic Press, San Diego, CA
Jolivet P (1998) Interrelationship between insects and plants.
CRC Press, Boca Raton, FL
House Dust Mite
Several species of mites in the family Pyroglyphi-
dae are considered to be dust mites.
 Mites
House Fly, Musca domestica L.
(Diptera: Muscidae)
John L. Capinera
University of Florida, Gainesville, FL, USA
This common fly originated on the steppes of
­central Asia, but now occurs on all inhabited
­continents, in all climates from tropical to temper-
ate, and in a variety of environments ranging from
rural to urban. It is commonly associated with
animal feces, but has adapted well to feeding on
garbage, so it is abundant almost anywhere people
live. It is the most common fly to invade homes,
and often is the dominant species around livestock
and poultry. Despite its inability to bite, it is both a
nuisance and a public health problem.
Life History
The life cycle of the house fly may be completed
in as little as 6 days, but under suboptimal con-
ditions may require up to 2 months. It breeds
continuously in warm climates such as in the
tropics and subtropics, producing more than 20
generations annually. Even in more temperate
areas they commonly undergo ten generations.
In cooler areas they overwinter as larvae or
House Fly, Musca domestica L. (Diptera: Muscidae)
H 1877
pupae, but adults perish when exposed to cold.
In heated areas in cold climates, however, they
breed throughout the winter months if the adults
have access to food and larvae to suitable devel-
opmental media.
The cylindrical-oval eggs (Fig. 54) are white,
and 1.0–1.2 mm long. Eggs typically are laid
in  clusters, often numbering 75–150 eggs per
cluster. A female normally deposits 2–6 clusters
of eggs during her life span. Maximum egg pro-
duction occurs at intermediate temperatures,
25–30°C. Often, several flies will deposit their
eggs in close proximity, leading to large masses
of larvae and pupae. Eggs must remain moist or
they will not hatch. Egg hatch usually occurs
within 8–20 h. Due to its rapid development time
and high reproductive capacity, the house fly has
the capacity to increase in abundance rapidly.
Indeed, scientists impressed with its reproduc-
tive capacity have ­calculated that, lacking mor-
tality, a single pair of  flies could grow to a
population of 191,010,000,000,000,000,000 in
only 5 months, enough to cover the entire earth
in a layer of flies several meters deep!
The larvae are creamy white, cylindrical,
legless, and taper to a point at the head. The
head of larval house flies, like most maggots,
have dark mouth hooks. There are three larval
instars. The optimal temperature for larval
development is 35–38°C, though larval survival
is greatest at 17–32°C. Larvae complete their
development in 4–13 days at optimal tempera-
tures, but require 14–30 days at temperatures of
12–17°C. The moisture level of the food affects
larval survival, and 50–70% moisture content
favors survival, though some survival can occur
at almost any moisture level. Nutrient-rich sub-
strates such as animal manure provide an excel-
lent ­developmental substrate. Very little manure
is needed for larval development, and sand or
soil ­containing small amounts of degraded
manure allows for ­successful belowground
development. At maturity, larvae are 7–12 mm
long, and prefer to ­disperse to a dry location to
pupate.
1878
H House Fly, Musca domestica L. (Diptera: Muscidae)
House Fly, Musca domestica L. (Diptera: M­ uscidae),
Figure 54  House fly stages of development:
above, adult; below (top to bottom), egg, larva and
pupa. Note that the eggs are normally deposited
in clusters. (Upper photo by Jim Castner, lower
by Lyle Buss, both of the University of Florida).
The mature larva pupates within the cuticle
of the last larval instar (this structure is called
the  puparium), though the shape of the pupa is
quite different from the larva, being bluntly
rounded at both ends. It is about 8 mm long. The
color of the pupa changes from yellowish, to red-
dish brown, to dark brown or black over the
course of the pupal stage. Pupae complete their
development in 2–6 days at 32–37°C, but require
17–27 days at about 14°C.
The adult is about 4–7.5 mm long. The female is
typically larger, and can be distinguished from the
male by the relatively wide space between the eyes
(in males, the eyes almost touch). The dull, grayish
fly bears four narrow dark stripes on the thorax. The
abdomen may be gray, but often is yellowish, and
has a narrow dark line dorsally. The wings are trans-
parent except for the dark veins. They have sponging
mouthparts, which allows them to lap up liquid
food. Thus, they cannot bite animals and humans.
Flies regurgitate readily, secreting saliva onto solid
foods so it can be liquefied and ingested. Adults may
live up to 2 months, but more typically live 2–3
weeks. Without food, they survive only 2–3 days.
Longevity is enhanced by availability of suitable
food, especially sugar. Access to animal manure does
not lengthen adult life and they live longer at cooler
temperatures. They require food before they will
copulate, and copulation is completed in as few as 2
min or as long as 15 min. Oviposition commences
4–20 days after copulation. Female flies need access
to suitable food (protein) to allow them to produce
eggs, and manure alone is not adequate. They prefer
sunlight, and are active fliers during warm days.
They are inactive at night, and commonly can be
seen perching on the ceilings of barns or other shel-
ters. Lacking buildings on which to perch, flies rest
on trees, poles, wires, and other elevated objects. If
adequate food and oviposition sites are available,
emerging flies generally remain in the area. Lacking
these conditions, however, flies will easily disperse a
kilometer or more in search of food.
According to a study conducted in Texas, USA,
breeding site suitability (in descending order), was
horse manure, human excrement, cow manure, fer-
menting vegetable, and kitchen waste. However,
another study found that structures ­containing
swine, horse, sheep, cattle, and poultry varied in fly
abundance, with swine facilities containing the most
and poultry the least. Fruit and vegetable cull piles,
partially incinerated garbage, and incompletely
composted manure also are highly favored sites for
breeding.
Damage
House flies are mechanical vectors of animal and
human pathogens, particularly enteric diseases. At

times, they have been found contaminated with
viruses, bacteria, fungi, protozoa, and nematodes.
These microbial organisms adhere to their tarsi, legs,
mouthparts, and elsewhere as the flies move about
their environment, and then these microbes can be
relocated when the flies move to new food sources.
Of particular concern is their movement from animal
or human feces to food that will be eaten uncooked
by humans. Also, when consumed by flies, some
pathogens can be harbored in the mouthparts or
­alimentary canal for several days, and then be trans-
mitted when flies defecate or regurgitate. In situations
where plumbing is lacking, such as open latrines,
serious health problems can develop, especially if
there are outdoor food markets, hospitals, or slaughter
houses nearby. Among the pathogens commonly
transmitted by house flies are Salmonella, Shigella,
Campylobacter, Escherichia, Enterococcus, Chlamydia,
and many other species that cause illness. These flies
are most commonly linked to outbreaks of diarrhea
and shigellosis, but also are implicated in transmission
of food poisoning, typhoid fever, dysentery, tuber­
culosis, anthrax, ophthalmia, and parasitic worms.
In addition to disease transmission, house
flies can be a severe nuisance. These flies alight on
and near people where they can be an irritant,
especially for people who cannot wave them away,
such as infants and the asleep or infirm. If left
undisturbed, the flies may feed on the secretions
of the mouth, nose and eyes. When human habita-
tions are in close proximity to animal production
facilities, the flies emanating from such operations
usually prove to be a point of contention, some-
times resulting in legal actions.
Management
Tolerance of flies depends greatly on circum-
stances. In sensitive environments such as food
preparation and packing facilities, restaurants, and
hospitals, even small numbers of flies cannot be
tolerated. In the context of livestock or poultry
production, however, some flies are inevitable.
Serious problems occur when cities or suburban
House Fly, Musca domestica L. (Diptera: Muscidae)
H 1879
development occur near poultry production facil-
ities, as residents usually will not tolerate the large
numbers of flies emanating from such facilities.
Fly populations are commonly monitored
directly using sticky tapes or ribbons, or baited
traps, to capture flies. Densities are also assessed
indirectly using spot cards to assess the number
vomit or fecal spots. Traps can be baited with
molasses, sugar, fruit or meat, and often are used
in combination with a device that captures the
attracted flies. The sex pheromone (Z)-9-tricosene
also functions as an aggregation pheromone, and
is called muscalure. Muscalure is formulated with
sugar as a commercially-available fly bait for local
population suppression, as well as an enhance-
ment for population monitoring.
Ultraviolet light traps can be used to assess
population levels, but also serve as a non-chemical
control technique that can be used indoors in both
agricultural and non-agricultural areas. They nor-
mally function by electrocuting flies that enter the
trap, though those used in restaurants typically
have a sticky panel. Flies do not orient to traps
from a great distance, so several are normally
needed for them to be effective. Placement should
include within 4–8 m of entryways, and within
1.5 m of the floor, to take advantage of fly flight
behavior. They should be operated continuously,
although they are most effective when the room
lights are off.
The most important approach for fly manage-
ment is sanitation. Flies should be deprived of
suitable oviposition sites, and larval environments
should be eliminated or made too dry for high lev-
els of survival to occur. To accomplish this, removal
of fecal material, agricultural refuse, spilled feed,
soiled animal bedding, and food wastes should
occur at least twice per week. Around homes and
businesses, screening or covering of windows,
doors or air doors, and trash containers proves
useful in denying access of flies to breeding sites.
Packaging household trash in plastic bags, and
burying trash under at least 15 cm of soil and in
sanitary landfills also helps to eliminate breeding.
Trash cans and dumpsters should have tight-fitting
1880
H Hover Flies
lids; failing this, slow release fumigant insecticide
dispensers are sometimes installed on the inside
of the lids to reduce fly survival.
In agricultural areas, manure can be scattered
over fields so that it quickly dries and becomes
unsuitable for egg and larval survival. Composting
of manure can be effective if the compost is prop-
erly maintained, including regular turning. Manure
can also be liquefied and stored in lagoons anaero-
bically, though at some point the solids need to be
separated. Manure can also be treated with an
insecticide, though this method is highly discour-
aged as it interferes with biological control of flies,
often resulting in a rebound of the fly population.
More commonly, insecticides (especially insect
growth regulators) can be fed to livestock, and
residual insecticide in the manure inhibits fly
breeding. In animal facilities, insecticides are often
applied to the favored resting places of adults, or
bait stations established to poison adults with either
solid or liquid formulations. Continuous exposure
of flies to insecticides has led to development of
insecticide resistance to many insecticides.
Natural biological suppression of the house fly
results primarily from the actions of certain chalci-
doid wasps (Hymenoptera: Pteromalidae), of which
many species have been associated with house fly
around the world. Among the more important are
Muscidifurax and Sphalangia spp. Ichneumonids
and other parasitoids, as well as some predatory
insects (especially histerids [Coleoptera: Histeri-
dae] and staphylinids [Coleoptera: Staphylinidae]),
also contribute to fly mortality, but under optimal
fly breeding conditions the house fly quickly
builds to high numbers. Augmentative biological
control using insectary-reared parasitoids has
been quite successful in some dairies, feedlots and
poultry house situations. The species most often
released for biological suppression in North
America are Muscidifurax raptor Girault and
Saunders, M. raptorellus Kogan and Legner,
­Spalangia endius Walker, and S. nigroaenea
Curtis. These different species function better
under different conditions, some performing
better under cooler or warmer conditions, others
parasitizing flies near the surface or deeper in the
pupation medium.
References
Barnard DR, Geden CJ (1993) Influence of larval density and
temperature in poultry manure on development of the
house fly (Diptera: Muscidae). Environ Entomol
22:971–977
Bishoff FC, Dove WE, Parman DC (1915) Notes on certain
points of economic importance in the biology of the
house fly. J Econ Entomol 8:54–71
Hogsette JA (1996) Development of house flies (Diptera: Mus-
cidae) in sand containing various amounts of manure
solids and moisture. J Econ Entomol 89:940–945
Howard LO, Bishopp FC (1925) The house fly and how to
suppress it. USDA Farmers’ Bulletin 1408, 18pp
Lysyk TJ (1991) Effects of temperature, food, and sucrose
feeding on longevity of the house fly (Diptera: Musci-
dae). Environ Entomol 20:1176–1180
Hedges SA (ed) (2004) The Mallis handbook of pest control,
9th edn. GIE Media, Cleveland. 1396 pp
Seymour RC, Campbell JB (1993) Predators and parasitoids
of house flies and stable flies (Diptera: Muscidae) in
cattle confinements in west central Nebraska. Environ
Entomol 22:212–219
Hover Flies
Members of the family Syrphidae (order Diptera).
 Flies
Howard, Leland Ossian
Leland Howard was born in Illinois on June 11,
1857, but moved with his parents to New York
state when he was young, and it was in the vicin-
ity of the town of Ithaca that he grew up. His
interests in natural history were encouraged by
his parents. However, when he entered Cornell
University, it was to study civil engineering at the
insistence of his mother, who by then was a
widow. Leland transferred to the natural sciences
without telling his mother, studied botany,

g­ eology, and chemistry, and worked in the labo-
ratory of Henry Comstock. He graduated in 1877,
and the following summer was contacted by
Charles Riley to persuade him to go to Washing-
ton, DC, and work for the U.S. Department of
Agriculture. The offer was accepted, and Howard
moved there and worked on numerous projects
in applied entomology, including brown-tail
moth, gypsy moth, European corn borer, Japa-
nese beetle, and scale insects, and taxonomy of
Hymenoptera including Ichneumonidae, Bra-
conidae, Chalcididae, and Proto­trupoidea. He
received an M.S. degree from Cornell University
for his thesis “The morphology of Chalcididae.”
He and Riley were among the founders of the
Entomological Society of Washington in 1884.
He married in 1886. In 1894 after the resignation
of Riley, Leland Howard was appointed Division
Chief in his place. Books followed, such as (1901)
“Mosquitoes, how they live, how they are classi-
fied, and how they may be destroyed,” (1901) “The
insect book,” (1911) “The housefly – disease car-
rier,” (1912–1917) with Dyar and Knab as coau-
thors “The mosquitoes of North and Central
America and the West Indies,” (1930) “A history
of applied entomology,” (1931) “The insect men-
ace,” and (1933) “Fighting the insects: the story of
an entomologist.” Awards followed in the form of
honorary doctoral degrees from Georgetown,
Pittsburgh, California, Toronto, and Rutgers uni-
versities, as well as an honorary M.D. degree from
Georgetown University. In 1894 he became presi-
dent of the American Association of Economic
Entomologists, and in 1898 permanent secretary
of the American Association for the Advance-
ment of Science. He retired in 1927 and died on
May 1, 1950, perhaps the most respected ento-
mologist in American history.
Reference
Mallis A (1971) Leland Ossian Howard. In: American ento-
mologists. Rutgers University Press, New Brunswick, NJ,
pp 79–86
Huffaker, Carl Barton
H 1881
Hübner, Jacob
Jacob Hübner was born in Augsburg, Germany,
on June 20, 1761. His early life seems unrecorded.
His great works were (1786–1790) “Beiträge
zur Geschichte europäischer Schmetterlinge,”
(1793) “Sammlung äuserlesener Vogel und
Schmetterlinge,” (1805–1824) “Sammlung euro­
päischer Schmetterlinge,” (1806–1818) “Geschichte
euro­päischer Schmetterlinge,” and (1806–1824)
“Sammlung exotische Schmetterlinge,” (1806)
“Tentamen determinationis, digestionis atque
denominationis singularum stirpium Lepidopter-
orum…,” and (1816) “Verzeichniss bekannter
Schmetterlinge.” All of the works deal with Lepido­
ptera (the second also with birds). Some of the
works are magnificently illustrated with copper
plates, making identification easy. The (1806–1824)
“Sammlung exotische Schmetterlinge” deals with
exotic species of Lepidoptera [meaning those that
occur abroad] and includes several North Ameri-
can species, some of which are pests. The (1806)
“Tentamen” was a work distributed to just a few
ento­mo­logists, in which the  author tried to fix
scientific names, whose validity was subsequently
hotly debated. He died in Augsburg on September
13, 1826. This man was the world’s first great
lepidopterist.
References
Essig EO (1931) Hübner, Jacob, p. 664–667 in A history of
entomology. Macmillan, New York, 1029 pp
Freyer CF (1861) Erinnerung an Jacob Hübner aus Augsburg.
Stettiner Entom Ztng 22:297–299
Huffaker, Carl Barton
Carl Huffaker was born in the state of Kentucky,
USA, on September 30, 1914. His B.S. (1938) and
M.S. (1939) degrees were from the University of
Kentucky, with a Ph.D. from Ohio State University
in 1942. His career began as medical entomologist
1882
H Human Botfly, Dermatobia hominis (Linneaus, Jr.) (Diptera: Qestridae)
with the U.S. Health and Sanitation Division of
Inter-American Affairs, in Colombia, Haiti, and
the Dominican Republic. He was recruited in 1946
by the Division of Biological Control of the Uni-
versity of California, stationed in Berkeley, and
remained in Berkeley for the rest of his career. His
work emphasized the biological control of weeds,
and a major one was control of Hypericum perfo-
ratum (perforated St. Johnswort, or Klamath
weed) in the Pacific coastal states of the USA.
He  published over 200 papers. In 1970–1983 he
was director of the University of California’s Inter-
national Center of Integrated and Biological Con-
trol. He was co-leader (with Paul DeBach) of the
­Integrated Pest Management Project, sponsored
by the U.S. Department of Agriculture, U.S. Envi-
ronmental Protection Agency, and National
­Science Foundation, involving scientists from 18
universities. This project was so much associated
with his leadership that it was widely known as
“The Huffaker Project.” He served as president of
the Entomological Society of America and was
elected to the National Academy of Sciences. He
died on October 10, 1995, after a prolonged illness,
and was survived by his wife and four children.
Reference
Caltagirone L, Dahlsten D, Garcia R, Gutierrez A, Hagen
K, Sylvester E (1995) Carl Barton Huffaker, Depart-
ment of Entomology and Parasitology, Berkeley.
Available at http://sunsite.berkeley.edu:2020/dynaweb/
teiproj/uchist/inmemoriam/inmemoriam 1995 Accessed
Aug 2002
Human Botfly, Dermatobia
hominis (Linneaus, Jr.) (Diptera:
Oestridae)
eugene j. gerberg
University of Florida, Gainesville, FL, USA
This large (approximately 12 mm) metallic blue fly
is also known as tropical warble fly, human warble
fly, and torsalo. The fly occurs in Mexico, Central
and South America from approximately 25°N to
32°S latitude. The fly, which does not feed, is found
primarily along the margins of forested areas. It
affects livestock, pests and other mammals in addi-
tion to humans. On occasion birds are affected, and
young livestock in Central and South America can
be so heavily infested that they are killed by this fly.
The life cycle is unusual, as the adult female botfly
(Fig. 55) lays its white eggs on day-flying flies, par-
ticularly mosquitoes, usually of the genus Psoro-
phora. The mosquito then inadvertently transports
the eggs of the botfly to distant vertebrate hosts. The
female lays 14–200 eggs on the ventral surface of
the transport insect, usually mosquitoes, but some-
times ticks serve as hosts. The eggs are 1 mm in
length. When the egg-bearing transport insect feeds
on a warm-blooded host, the heat stimulates the
Dermatobia eggs to hatch, the larva emerges and
bores into the skin. They usually penetrate at the
feeding site of the mosquito, but also can penetrate
unbroken skin at the hair pore. This botfly has also
been known to lay eggs on wet laundry. When a
person puts on the egg-infested clothes, the larvae
emerge from the eggs and bore into the skin. Within
the skin, the larva grows, molts, and forms a cyst,
boil or warble. In humans, lesions develop mainly in
Human Botfly, Dermatobia hominis (Linneaus, Jr.)
(Diptera: Oestridae), Figure 55  Adult female
human botfly, Dermatobia hominis (adapted from
James 1947).

the hands, wrists, ankles, neck, face and head, but
occasionally also in the brain. Several larvae can
inhabit the host simultaneously, and they normally
stay close to the site of entry, using the entry pore as
a breathing site. The inflamed pocket may itch, lar-
val movement is painful, and eventually discharge
emanates from the swelling. Larval development
time is about 1–4 months. Though not generally
considered life-threatening, it can be fatal to very
young children (<5 years of age). The adult lifespan
is brief, usually only a few days.
Treatment of humans usually involves exer-
cising pressure on or around the lesion, resulting
in expulsion. This may not be effective, however,
and excision follow by treatment to prevent sec-
ondary infection may be preferable. Secondary
infection is common.
The larva has twelve body segments. The first
segment has mouthparts with hooks. Most of the
Human Botfly, Dermatobia hominis (Linneaus, Jr.)
(Diptera: Oestridae), Figure 56  Mature larva of
human botfly, Dermatobia hominis, shown
infesting its host, with the posterior spiracles
protruding from a hole in the host’s skin (adapted
from Craig and Faust 1940).
Human Lice
H 1883
body segments have spines. The larva pushes its
posterior spiracles through the aperture in the skin
to maintain a hole open to the external air. After
several weeks the mature larva (Fig.  56) emerges
during the night or early morning, and drops to the
ground to pupate. The larval period lasts approxi-
mately 40 days. The last larval skin serves as a shell
for the pupa. The pupal period lasts from 14 to 24
days, and then the adult emerges. The emerging
females mate and seek a transport insect.
Management includes using mosquito
­repellents on the skin and clothing, and mosquito
netting around beds. This benefits not only
human botfly management, but helps prevent
mosquito-vectored diseases such as dengue, yel-
low fever, and malaria.
References
Bates M (1943) Mosquitoes as vectors of Dermatobia in eastern
Colombia. Ann Entomol Soc Am 36:21–24
Banegas AD, Mourier H, Graham OH (1967) Laboratory
colonization of Dermatobia hominis (Diptera: Cutereb-
ridae). Ann Entomol Soc Am 60:511–514
Catts EP (1982) Biology of New World bot flies: Cuterebridae.
Ann Rev Entomol 27:313–318
Guimarães JH, Papavero N (1966) A tentative annotated
bibliography of Dermatobia hominis (Linnaeus Jr. 1781)
(Diptera, Cuterebridae). Arquivos de Zoologia do
Estado de São Paulo 14:223–294
Koone HD, Banegas AD (1959) Biology and control of
Dermatobia hominis (L. Jr.) in Honduras. J Kansas Entomol
Soc 32:100–108
Rosen IJ, Neuberger N (1977) Myiasis Dermatobia hominis
Linn: report of a case and review of literature. Cutis
19:63–66
Sancho E (1988) Dermatobia, the neotropical warble fly.
Parasitol Today 4:242–246
Human Lice
terri l. meinking
University of Miami School of Medicine, Miami,
FL, USA
Lice are wingless, six-legged, bloodsucking
insects that belong to the order Siphunculata
1884
H Human Lice

(also called Anoplura, or combined with the
chewing lice and called Phthiraptera). There are
nearly 4,000 species of lice recognized, although
only 560 species suck blood and feed on mam-
mals. Lice are very host specific; therefore,
human lice cannot be transmitted between mam-
mals. There are only three species of lice that
infest humans: Pediculus humanus humanus, the
body or “clothing” louse; Pediculus humanus
capitis, the head louse; and Phthirus (or Pthirus)
pubis, the pubic or “crab” louse (Fig. 57).
Biology
Lice are hemimetabolic, meaning that they do
not go through a complete metamorphosis like
mosquitoes or fleas. After a 7–12 day incubation
period, nymphs are hatched from eggs or “nits.”
Almost immediately, the nymph must feed on a
human host to avoid death from starvation and
dehydration.
Nymphs resemble miniature adults and
undergo three nymphal instars before maturation.
Several feedings occur between each molt of the
­chitinous exoskeleton. When feeding, the body
structure of the louse is capable of expanding by one
third of its body weight per meal. Crab and head lice
feed regularly every 4–6 h, although the body louse
can survive several days without a blood meal.
It is not until the 3rd and final molt that the
sex is determined. It is more difficult to identify
the gender of P(h)thirus than Pediculus. Within 2
days of maturation, the female will feed several
times, copulate, and begin laying eggs. The female
head louse lays an average of 3–6 eggs per day,
with crab lice laying fewer and body lice laying
more. The female louse attaches nits to hairs

Human Lice, Figure 57  Human lice. Upper left: adult female head louse with blood meal; upper right:
alternating male and female head lice (male-female-male-female); lower left: nymph of crab louse; lower
right: scanning electron micrograph of head louse egg (nit).

(or  ibers in the case of body lice) by secreting a
glue that hardens on contact with air.
In each species, females are usually 20% larger
than males, with longer, wider, and rounder bodies.
Only males have dark brown stripes on their
back. Only females, however, have an invaginated
V-shape located on the posterior end of their body,
in order to deposit eggs on the hair shaft or cloth
fiber. The average life span of a louse is 30–42 days.
Habitat and Epidemiology
Pediculus humanus
The body louse does not live on the body, but in
clothing or bedding, and only travels to the host to
feed. Nits are laid on cloth fibers, particularly in
the seams and collars. Although people associate
all lice with poor hygiene, only body lice infesta-
tions are due to lack of cleanliness. Infestations are
common in individuals who are unable to change
or wash their clothing. Frequently, these people
are forced to live in crowded, unsanitary environ-
ments, such as those exposed to war, natural disaster,
refugee status, or homelessness.
Pediculus capitis
Head lice are 25% smaller than body lice, but other-
wise are identical in appearance. Head lice live on
the scalp, preferring a clean, healthy head. Nits are
usually laid close to the scalp for warmth, although
in tropical climates, they can be found anywhere on
the hair shaft. The female louse deposits nits on the
hair shaft with a biological adhesive for which there
is no known solvent. Hair debris such as gel, spray,
and dandruff, are often mistaken for nits. However,
hair residue slides easily off the hair, whereas nits
are firmly cemented to the hair shaft. Head lice
infestations are most common in children aged
3–11 due to their frequent head to head contact
during play. Other close contacts, such as family
members, teachers and day care workers, may also
Human Lice
H 1885
become infested. Head lice transmission can also
occur by sharing of brushes, combs, hats, helmets,
and other headgear and hair accessories.
P(h)thirus pubis
Crab lice are not discriminatory in host choice, and
are found in all levels of society and various ethnic
groups. Crab lice are typically located in the pubic
and perianal areas, but can be also found on any
hairy part of the body. Secondary sites of infesta-
tion commonly include beards, mustaches, scalp,
eyebrows, eyelashes, and axillae. Transmission
­generally occurs by sexual contact, although crab
lice can also be transmitted by infested towels and
bedding. It is not uncommon to find crab lice with
their craws caught in the fiber loops of a towel
recently used by an infested individual. Fomite
transmission occurs more commonly than origi-
nally thought because adult crab lice are viable off
the host for at least 36 h.
Disease Transmission
Pediculus humanus
Disease transmission is not caused by the bite of
the louse, but by infected fecal pellets that can be
scratched into the bite site or by an abrasion in
the skin. The body louse is known to transmit the
­following diseases: epidemic typhus (caused by
­Rickettsia prowazekii), murine typhus (caused by Rick-
ettsia typhii, trench fever (caused by ­­Bartonella
quintana), and relapsing fever (caused by Borrelia
recurrentis).
Pediculus capitis and P(h)thirus pubis
Head and crab lice are known to mechanically trans-
mit group A Streptococcus pyogenes and Staphylo-
coccus aureus. Head and crab lice have not yet been
investigated as transmitters of blood borne diseases.
1886
H Humeral
Treatment
Pediculus humanus
Treatment of body lice includes washing clothing
and bedding in hot water and then drying in a
hot cycle (65°C, 149°F). Drug therapies, such as
permethrin or malathion dusting powders, are
effective treatment methods in instances of mass
infestation. In individual cases, prescription per-
methrin 5% topical cream or oral ivermectin are
recommended treatments.
Pediculus capitis
Treatments for head lice include lindane, mala-
thion, permethrin or natural pyrethrin products.
Lindane products demonstrate resistance and
may have harmful toxicity effects to the central
nervous system (CNS). Over-the-counter treat-
ment with 1% permethrin or natural pyrethrin
products can be used, but resistance has been
reported in the United States and other countries.
Currently, the most effective treatments for head
lice include prescription malathion lotion, oral
ivermectin (off-label), certain products with anise
oil and alcohol, and nit combs as adjunct therapy.
P(h)thirus pubis
Although head lice products may be used in
treating crab lice, 5% permethrin cream (pre-
scription treatment for scabies) or oral ivermec-
tin are the most effective treatments. Medication
should be applied to all hairy areas of the body,
including the scalp.
Phthiriasis palpebrarum
Topical treatments should not be used around
the eye area. Vaseline is an effective treatment for
crab lice of the eyelashes, as it suffocates lice and
eggs and lubricates the lashes, making nit removal
easier and is non-toxic.
 Chewing and Sucking Lice
 School IPM
References
Alexander JO (1984) Arthropods and human skin. Springer-
Verlag, Berlin, Germany
Andrews M (1976) The life that lives on man. Taplinger, New
York, NY
Burgess I (1995) Human lice and their management. Adv
Parasitol 36:271–342
Chosidow O (2000) Scabies and pediculosis. Lancet 355:819–826
Meinking TL, Taplin D (1995) Infestations. In: Schachner LA,
Hansen RC (eds) Pediatric dermatology, 2nd edn.
Churchill Livingstone, New York, NY, pp 1347–1392
Meinking TL (1999) Infestations. Curr Prob Dermatol
11:75–118
Mumcuoglu KY, Hemingway J, Miller J, Ioffe-Uspensky I,
Klaus S, Ben-Ishai F, Galun R (1995) Permethrin resis-
tance in the head louse Pediculus humanus capitis from
Israel. Med Vet Entomol 9:427–432
Humeral
Pertaining to the “shoulder” (near the point of
attachment of the wing) of the insect, or located in
the anterior basal portion of the wing.
Humeral Angle
The angle of the base of the costal margin of the
wing, near the point of attachment.
 Wings of Insects
Humeral Cross Vein
The cross vein extending between the costa and
the subcosta near the base of the wing.
 Wings of Insects
Hummingbird Lice
Members of the family Ricinide (order
Phthiraptera).
 Chewing and Sucking Lice

Hummingbird Moths
Some members of the family Sphingidae (order
Lepidoptera).
 Hawk Moths
 Butterflies and Moths
Humpbacked Flies
Members of the family Phoridae (order Diptera).
 Flies
Human Lymphatic Filariasis
(Elephantiasis)
jai k. nayar
University of Florida, Vero Beach, FL, USA
Human lymphatic filariasis (elephantiasis) is a
debilitating and deforming disease caused by
infection with parasitic round worms belonging to
the phylum Nematoda, order Spirurida, superfam-
ily Filarioidea, and family Filariidae, genera
Wuchereria and Brugia, species W. bancrofti and
species B. malayi and B. timori. Humans are the
final and exclusive host of W. bancrofti. Brugia
malayi, in addition to infecting humans, has other
strains that infect some feline and/or monkey spe-
cies, but the life-cycles in humans and in these
other animals generally remain epidemiologically
distinct. These parasites complete their develop-
ment in two host types, the human (the final
and definitive host) or other vertebrates, and
the mosquito (the intermediate host). The disease
spreads from person to person by the bites of
infected mosquitoes.
Approximately 120 million people are infected
in at least 80 countries of the world throughout
the tropics or subtropics (41 North latitude to 31
South latitude), that include South America, Cuba,
Puerto Rico, West Indies, Africa, Spain, Turkey, Asia,
Australia, and many South Pacific Islands. It is
­estimated that one billion (20% of the world’s
Human Lymphatic Filariasis (Elephantiasis)
H 1887
­ opulation) are at risk of acquiring infection.
p
Ninety percent of these infections are caused by
W. bancrofti. The remaining 10% of infections
caused by B. malayi are restricted to southeast
Asia, including India, Thailand, Vietnam, southern
China, and the south Pacific islands, and by B. timori
in the Lesser Sunda Islands of eastern Indonesia.
Both W. bancrofti and B. malayi have four
types based on the density of microfilariae in the
peripheral circulation: nocturnally periodic, noc-
turnally subperiodic, diurnally subperiodic and
diurnally periodic. Nocturnally periodic type of
W. bancrofti is generally found in the slum areas of
cities (urban type) and rural areas (rural type). A
nocturnally subperiodic type is found in Thailand
and Vietnam. The diurnally subperiodic type is
rural and found in the eastern Pacific islands and
the diurnally periodic type of B. malayi is mostly
found in areas with irrigated rice fields.
The adult worms that cause the disease live for
7–10 years in the victim’s lymph vessels, near to and
in lymph nodes. Adult females are long and slender
(W. bancrofti – 6–10 cm long and 300 μm wide,
B. malayi – 48 mm long, and B. timori – 40 mm
long) with a smooth cuticle and bluntly rounded
ends. The head is slightly swollen and bears two
circles of well-defined papillae. The mouth is
small;  a  buccal cavity is lacking. Males are much
smaller (mean length for W. bancrofti – 40 mm,
B. malayi – 18 mm, and B. timori – 15 μm) with a
fingerlike tail. The vulva of the female is near the
level of the middle of the esophagus. Adult females,
after mating, produce microfilariae (mean length
for W. bancrofti – 290 μm, B. malayi – 222 μm and
B. timori – 310 μm) which are found in the blood
and peripheral circulation. In endemic areas, in the
nocturnally periodic type, the level of microfilariae
in the peripheral circulation of humans is controlled
by a circadian periodicity or 24 h light-dark cycle,
that is, the numbers of microfilariae in the periph-
eral circulation show a marked peak during the
night. In the day time the microfilariae move mainly
to the pulmonary capillaries and very few are found
in the peripheral circulation. The appearance of
large numbers of microfilariae in the peripheral
1888
H Human Lymphatic Filariasis (Elephantiasis)
c­ irculation at night coincides with the biting activ-
ity of the local mosquito vector. In the nocturnally
­subperiodic, diurnally subperiodic and diurnally
periodic types, microfilariae can be found in the
peripheral circulation at all hours of day and night.
The microfilariae do not develop further in humans,
but only in susceptible species of mosquitoes.
Development of microfilariae to the L3 or
infective larval stage in mosquitoes is identical in
all three species (Fig. 58). In mosquitoes, ingested
microfilariae in the blood meal do not multiply
but grow to the L3 stage after two molts with
2 weeks. Within a day after the microfilariae are
ingested, they shed their sheath and penetrate the
midgut wall, move in to the hemocoel, then
migrate to the thoracic muscles and lodge them-
selves intracellularly. The slender active microfi-
laria transforms to the short thick inactive
sausage-stage or L1 larva within the next 2–4
days. The L1 larva has a cuticle which forms a
conspicuous tail, characteristic of this stage. In
the genus Brugia one to two nuclei are present in
the tail. Five to six days after being ingested as
microfilariae, the L1 larva molts to the L2 stage.
The L2 larva grows rapidly and has a thin cuticle
which can be seen at the caudal end as a short
tail. There are one or two papillae at the caudal
end. During the second molt the larva sheds its
cuticle and becomes the infective L3 larva. The L3
larva grows further in length but not in width,
moving actively to the hemocoel of the mosquito,
first towards the abdomen and later to the head
and proboscis. The caudal end of W. bancrofti is
characterized by having three teat-like papillae of
equal size, whereas in the Brugia, the central
papilla is the most prominent. The duration of lar-
val development in mosquito vectors is affected
by the ambient temperature; generally, the
warmer it is the more rapid the development. It
usually takes 10–14 days for W. bancrofti to reach
the infective stage, and 7–10 days for Brugia species.
When the now infective mosquito bites a
human, some or all the infective larvae escape
from the proboscis onto the skin and actively enter
the wound made by the mosquito. The L3 larvae
move to the lymphatic system and start to develop
and molt to the L4 stage followed by the L5 or
young adults and finally to the mature male and
female worms. After fertilization, the female
worms produce microfilariae, which find their way
from the lymphatic system to the blood circula-
tion. The pre-patent period from the entrance of
L3 when the mosquito bites the human to the
appearance of microfilariae in the peripheral
blood, is about 3–4 months for Brugia sp. and
about 8–9 months for W. bancrofti.
The intermediate hosts (mosquitoes belong-
ing to the genera Aedes, Culex, Anopheles, and
Mansonia – about 77 species) become infected
when they take an infective blood meal from an
infected definitive host (man). Urban W. bancrofti
with noctural periodicity is transmitted mainly by
Culex quinquefasciatus in tropical regions, by
Culex pipiens pallens in China and Japan, and
Culex pipiens molestus in the eastern Mediterra-
nean. Rural W. bancrofti that is nocturnally subpe-
riodic over most of its range is mainly transmitted
by several species of Anopheles in the western
Pacific islands and Africa, occasionally Aedes spp.
in eastern Pacific islands, and rarely by Mansonia
uniformis in Sri Lanka. The diurnally subperiodic
type is transmitted predominantly by several
species of Aedes, and the nocturnally subperiodic
type by the mosquitoes of the Aedes niveus group.
The nocturnally periodic type of B. malayi is trans-
mitted in certain regions by Mansonia spp. and in
other regions by Anopheles barbirostris or An.
campestris in Indonesia and Malaysia. Aedes togoi
is a vector in coastal parts of the Republic of Korea,
and parts of southern China. The nocturnally
subperiodic type of B. malayi is transmitted by
Mansonia spp. B. timori is transmitted by Anopheles
barbirostris.
Clinical course of lymphatic filariasis in
humans can be divided into three distinct stages,
asymptomatic, acute and chronic, generally pro-
gressing in that order. The asymptomatic stage is
characterized by the presence of microfilariae in
the peripheral blood, although there are no clinical
manifestations of filariasis. The acute symptomatic
 Human Lymphatic Filariasis (Elephantiasis)
H 1889

Life cycle of lymphatic filariasis

L3 and L4 larvae develop

to adult worms in skin Adult worms
and lymphatic channels

Final host
Infective larvae (L3)
Woman with filariasis
enter mammalian skin
through wound made
Mf. in blood
by proboscis of mosquito
circulation

Mf. in midgut of
female mosquito
Infective larvae (L3)
coming from proboscis
L1
Thoracic
Mf. muscles L2
Midgut

L3

L3 Lm
Mf.
Intermediate host
Developmental stages Lb Hphy
in mosquito

Human Lymphatic Filariasis (Elephantiasis), Figure 58  Life cycle of lymphatic filariasis. Stages of filarial
worm in human: L3 = third stage larvae and L4 = fourth stage larvae. Stages of filarial larvae in the
mosquito: mf = microfilariae, L1 = first stage larvae (sausage stage), L2 = second stage larvae, and
L3 = third stage larvae. Larval stages L1 to L3 are shown both as normal larvae and in histological sections.
Lm = labium, Lb = labrum, and Hphy = hypopharynx.

stage is characterized by recurrent attacks of fever
associated with inflammation of the lymph nodes
(lymphadenitis) and lymph vessels (lymphangitis).
In bancroftian filariasis recurrent attacks of fever
associated with lymphadenitis are less frequently
seen than in brugian filariasis. In addition to the
lymph nodes in the inguinal, axillary and epitro-
chlear regions, the lymphatic system of the male
genitalia is frequently affected. Typically, each
attack of fever and lymphadenitis lasts for several
days or weeks and usually subsides spontaneously
following bed-rest. The chronic signs of filariasis
do not develop usually before the age of 15 years
and only a small portion of the community is
affected. During the chronic stage microfilariae are
usually absent from the blood.
1890
H Human Scabies
In bancroftian filariasis, the occurrence of
the major chronic signs are accumulation of
serous fluid in the body cavity (hydrocele), appear-
ance of microfilariae in urine (chyluria), swelling
of body parts (lymphodema) and hardening and
thickening of the skin of swollen body parts (ele-
phantiasis), which may differ from one area to
another. The most common are hydrocele and
swelling of the testis, followed by elephantiasis of
the entire lower limb, the entire arm, the vulva,
and the breast, in descending order of frequency.
In brugian filariasis, the leg below the knee or the
arm below the elbow are characteristically affected.
Genital involvement has not been reported for
brugian filariasis.
The worms in the lymphatic system cause tis-
sue changes which restrict normal flow of lymph
and result in swelling, fibrosis and eventually sec-
ondary infections in the affected tissues. The lower
extremities and groin are the parts most likely to
be affected. In its severest form, lymphatic filariasis
causes elephantiasis, or dramatic swelling of limbs
(usually the leg) and genitals (usually the scro-
tum). These conditions have a devastating effect
on the quality of life of those affected, impacting
them not only physically, but also emotionally and
economically.
Human lymphatic filariasis is a disease which
develops over several years and is localized in
communities. Individuals can be treated with
albendazole plus either diethylcarbamazine (DEC)
or ivermectin (Mectizan®), which have been
shown to be effective in removing microfilariae
from the blood for a full year after treatment and
slowly killing the adult stage of the parasite.
Recently, the World Health Organization has
adopted the policy of a Global Program to Elimi-
nate Lymphatic Filariasis by 2020. The strategy for
such a program is based on two concepts: first, to
interrupt transmission of infection and second, to
alleviate the suffering of infected individuals
through management of the disease. The first con-
cept includes treatment of each individual in the
community with a single dose of either of the two-
drug combinations, albendazole and DEC or
albendazole and ivermectin, once a year for sev-
eral years and to implement vector control prac-
tices in each infected community. The second
concept is to alleviate the suffering caused by the
disease by educating the community about the
programs to raise awareness in affected patients.
This would promote the benefits of intensive local
hygiene, vector control and possible improvement,
both in the damage that has already occurred, and
in preventing the debilitating and painful, acute
episodes of inflammation.
References
Anon (1987) Control of lymphatic filariasis. A manual for
health personnel. World Health Organization, Geneva,
Switzerland, 89 pp
Anonymous (1992) Lymphatic filariasis: the disease and its
control. Fifth report of the WHO Expert Committee on
Filariasis. WHO Technical Report Series No. 821. World
Health Organization, Geneva, Switzerland
Anonymous (1999) Lymphatic filariasis. WHO information
fact sheet. World Health Organization, Geneva, Switzerland,
3 pp
Anonymous (1999) Global program to eliminate lymphatic
filariasis. An informal report from WHO/CDS/CPE/
CEE/Fil. World Health Organization, Geneva,
Switzerland
Sasa M (1976) Human filariasis. University of Tokyo Press,
Tokyo, Japan
Human Scabies
john p. smith
Florida A & M University, Panama City, FL, USA
Scabies is a skin condition, also known as “7 year
itch” or “Norwegian itch,” produced by almost
invisible parasitic mites commonly referred to as
“itch mites,” Sarcoptes scabiei (DeGeer).
There are several varieties of itch mites that
can be distinguished only by the host they attack.
Dog, pig, horse and human itch mites are known
to exist. Though it is possible for people to be
infested with non-human species, usually little to
no adverse effect is produced.

Biology
Itch mite nymphs and adults are whitish in color
and have eight short, pointed legs equipped with
suction devices on the two front pairs and long
hairs on the remaining hind legs (Fig. 59). The life
cycle of the human itch mite consists of the egg,
six-legged larval, eight-legged nymphal and adult
stages. The period from egg to mature adult takes
10–14 days at body temperature. Itch mites seldom
survive more than a few hours off the host.
Injury and Spread
Mature female mites are responsible for the major-
ity of the skin irritation caused by tunneling in the
upper skin layer. Eggs are deposited in these tun-
nels that extend over one inch in length at 2–3 day
intervals over a 2-month period. Once the eggs
hatch in 3–8 days, emerging larvae exit the tunnels
and remain on or near the skin surface until they
reach adult stage. During this time, male mites
move about the surface where it is thought they
mate with females. These newly fertilized female
Human Scabies, Figure 59  The human itch mite,
Sarcoptes scabiei (actual size about the size of
this period “.”).
Human Scabies
H 1891
mites are highly active and are thought to be the
contagious stage. Itch mites are primarily transmit-
ted through close personal contact, though bed lin-
ens and clothing may serve as secondary sources.
Signs, Symptoms and Diagnosis
Red patches of skin on various regions of the body
characterize scabies. The most common infested
areas include: wrists, elbows, breasts, penis, bends
of the knees, between the fingers, and between the
buttocks. Sometimes the actual raised tunnels or
bumps produced by the mites can be seen. Tunnel-
ing mites may release toxic substances that pro-
duce severe itching. Often, constant itching
causes the individual to become pale and haggard
from loss of sleep. Scratching to relieve itching fre-
quently leads to secondary bacterial infections
that may become more serious than actual scabies.
Widespread allergic reactions also may occur due
to the mite presence.
First-time infestations often cause no immedi-
ate itching, although sensitivity normally occurs
after 1 month. Thereafter, subsequent infestations
result in reactions within the first 24 h. Infestations
producing these symptoms typically involve less
than 12 adult mites. Diagnosis is best made by apply-
ing mineral oil to the affected skin surface where
tunnels occur, scraping the skin with a scalpel, and
examining for the mites under a microscope.
Control
Recognize signs and symptoms of a possible
infestation and attempt to recover mites for
identification.
Inspect all individuals coming in contact with
infested person(s).
Treat all infested persons with one of the rec-
ommended miticides listed in the following treat-
ment section.Prophylactic treatment of non-infested
persons should be avoided unless screening pro-
grams are not effectively suppressing an outbreak.
1892
H Hungerford, Herbert Barker
Isolate infested person(s) for 24 h after treat-
ment has been applied.
Machine wash bed linens, clothing and all
other possible contaminated articles in hot (130°F)
water and soap, or tightly seal non-washables in
plastic bags for 2 weeks.
Treatment
Ointments containing 5% permethrin cream (Elim-
ite®), benzyl benzoate, malathion (Derbac-M®), cro-
tamiton (Eurox®), tetraethylthiuram monosulfide
(Tetmosol®), sulfur, thiabendazole, or 1% lin-
dane (Kwell®) are the most widely recommended
medications for itch mite control. These products
are available only through a physician by prescrip-
tion and should be applied strictly according to
labeled and/or prescribed directions.
 Mites
References
American Academy of Dermatology (1999) Available at
http://www.aad.org/education/students/parainfest.htm
Accessed January 2008
Benenson AS (ed) (1990) Scabies. Control of communicable
diseases in man, 15th edn. American Public Health
Association, Washington, DC, pp 385–388
Centers for Disease Control and Prevention (1999) Parasitic
diseases. Scabies. http://www.cdc.gov/ncidod/dpd/para
sites/scabies/default.htm
Indian and Inuit Health Committee, Canadian Pediatric Soci-
ety (1994) Scabies management. Can J Paediatr
1:152–155
Greene A (1996) What is the appropriate medication to use
for scabies? Available at http://www.drgreene.com/
961118.html
Hungerford, Herbert Barker
Herbert Hungerford was born in Kansas on
August 30, 1885. He grew up on a farm and
trained as a teacher, becoming teacher, principal,
and superintendent of schools between 1904 and
1909. Having married in 1905, he entered the
University of Kansas in 1909 and obtained a
bachelor’s degree in 1911. He immediately
became a member of the university faculty, and
in 1924 was appointed head of the Department
of Entomology and State Entomologist. His
research interest was the taxonomy, ecology and
behavior of water bugs, especially Corixidae and
Notonectidae. He built a large collection of
aquatic and semiaquatic Hemiptera on which he
continued working after retirement in 1956. In
1936, he was elected president of the Entomo-
logical Society of America and in 1953 of the
Society of Systematic Zoology. He died on May
13, 1963.
References
Mallis A (1971) Herbert Barker Hungerford, In: American
entomologists. Rutgers University Press, New Brunswick,
NJ, pp 236–238
Woodruff LC (1963) Obituary of Herbert Barker Hungerford.
J Kansas Entomol Soc 36:197–199
Hunting Billbug, Sphenophorus
venatus vestitus Chittenden
(Coleoptera: Curculionidae)
This billbug has become a serious problem for
some turfgrasses.
 Turfgrass Insects and their Management
Hyaline
Transparent, colorless, or glass-like.
Hyblaeidae
A family of moths (order Lepidoptera). They com-
monly are known as teak moths.
 Teak Moths
 Butterflies and Moths
 Hygrokinesis
H 1893

Hybosorid Scarab Beetles Hydrophilidae

Members of the family Hybosoridae (order A family of beetles (order Coleoptera). They com-
Coleoptera). monly are known as water scavenger beetles.
 Beetles  Beetles

Hybosoridae Hydropsychidae
A family of beetles (order Coleoptera). They com- A family of caddisflies (order Trichoptera). They
monly are known as hybosorid scarab beetles. commonly are known as net-spinning caddisflies.
 Beetles  Caddisflies

Hybothiridae Hydropyle
A family of sucking lice (order Phthiraptera).
Hydropyles are structures that allow the uptake of
 Chewing and Sucking Lice
water, and are found on some eggs. Water is
absorbed when the embryo is rapidly growing,
Hybrid Vigor and the egg enlarges in size. The eggs of most
aquatic insects absorb water, but terrestrial insects
Increased vigor resulting from the crossbreed- that deposit their eggs where moisture may occur
ing of two genetic lines (races, strains). The off- (e.g., grasshopper eggs in soil) also may have eggs
spring of such crossbreeding are more virgorous that enlarge. The hydropyle is formed interiorly
than either parent. from the serosal layer, but may be visible externally
by an extension of the chorion.
 Eggs of Insects
Hydraenidae
A family of beetles (order Coleoptera). They com-
monly are known as minute moss beetles.
Hydroscaphidae
 Beetles
A family of beetles (order Coleoptera). They com-
monly are known as skiff beetles.
Hydrobiosidae  Beetles

A family of caddisflies (order Trichoptera). They

commonly are known as microcaddisflies. Hydrostatic Skeleton
 Caddisflies
Maintenance of body form in soft-bodied insects
such as caterpillars by muscles exerting pressure
Hydrometridae on the fluid-filled body.
A family of bugs (order Hemiptera). They some-
times are called water measurers or marsh Hygrokinesis
treaders.
 Bugs A kinesis response with respect to moisture.
1894
H Hygrotaxis
Hygrotaxis
A taxis response with respect to moisture.
Hymenopodidae
A family of praying mantids (Mantodea).
 Praying Mantids
Hymenoptera
An order of insects. They commonly are known as
wasps, ants, bees and sawflies.
 Wasps, Ants, Bees and Sawflies
Hyocephalidae
A family of bugs (order Hemiptera, suborder
Pentamorpha).
 Bugs
Hypericin
Cyrus Abivardi
Swiss Federal Institute of Technology, Zurich,
Switzerland
Hypericin, well known for its photodynamic
action, is an aromatic polycyclic compound (a
highly condensed quinone) mainly synthesized by
plants of the genus Hypericum (family Hyperi-
caceae). A survey covering 200 species within the
family Hypericaceae, revealed that ca. 60% of these
plants contained Hypericin. It is usually confined
to certain species of the genus Hypericum, e.g., the
St. John’s worts, a common pasture weed through-
out the world. While this compound is found in
the leaves, stem and flowers of Hypericum perfora-
tum, it occurs only in multicellular trichomes of
the calyx of H. hirsutum.
Hypericin (Fig.  60) and its derivatives have
also been recorded from other sources. Hypericin
has been found in the chromophore of the photo-
receptor of Stenor coeruleus (a blue-green ciliate)
and in the integument of an Australian insect
(Nipaecoccus aurilanatus). Fagopyrin (a derivative
of hypericin) is found in buckwheat (Fagopyrum
esculentum). In addition, pencilliopsin, which
can be oxidized and irradiated to form hypericin,
has been isolated from a mold (Penicilliopsis
clavariaeformis).
Biological Activity
Effects on Insects
Although photodynamic reactions are generally
not species specific, the phototoxicity of hypericin
is known in only a few systems. Its phototoxicity
toward grazing animals, antiviral activity against
HIV-1, and insecticidal effect on Aedes mosquito
larvae have been reported.
It is well established that the active wave-
lengths in hypericin toxicity are greater than
500 nm. For larvae of the sphingid Manduca
sexta, the LD50 of purified hypericin was 16 μg/g
larval initial fresh weight in constant light (22
W/m2), but reduced irradiance resulted in
decreased mortality. Sublethal applications of
OH O OH
HO CH3
HO CH3
OH O OH
Hypericin, Figure 60  Structure of hypericin
(C30H16O8). an aromatic polycyclic compound
­mainly synthesized by plants of the genus
­Hypericum (family Hypericaceae).

hypericin retarded larval growth (body fresh
weight) in a dose-dependent manner. While
the phototoxic effect was lost when larvae were
maintained in darkness for more than 8 h before
irradiation, the potential for light-dependent
mortality was retained if larvae were starved
before irradiation.
Interestingly, several insects with different life-
styles have been reported to overcome the photo-
toxic effects of hypericin by appropriate behavioral
and biochemical strategies: (i) by avoiding light,
(ii) by feeding on the plant parts lacking hypericin,
or (iii) by decomposing hypericin. Studies on the
behavioral and biochemical adaptations of gener-
alist and specialist herbivorous insects feeding on
Hypericum perforatum (Guttiferae) have revealed
that the generalists Tettigonia viridissima L., Ruspo-
lia nitidula Scopoli, and Conocephalus discolor
Thunberg preferentially fed on the part of the
leaf lacking the phototoxic, hypericin-laden dark
glands. In contrast, the specialists Galeruca tana-
ceti L., Chrysolina geminata Paykull, and Cloantha
perspicillaris Boisduval showed no discriminatory
feeding pattern but exhibited a negative phototaxis
that is presumed to be an efficient strategy to over-
come the light-induced toxicity of hypericin.
Another mechanism of defense for insects
feeding on phototoxic plants may be the presence
of antioxidant enzymes such as superoxide dis-
mutase (SOD), catalase (CAT), glutathione per-
oxidase (GPOX), and glutathione reductase (GR).
The activities of these enzymes were examined in
the larvae of three lepidoptera: Ostrinia nubilalis,
Manduca sexta, and Anaitis plagiata. The highest
levels of antioxidant enzyme activity were found
in A. plagiata, a specialist feeder on Hypericum
perforatum, a plant which contains high levels of
the phototoxin hypericin. Larvae of A. plagiata
that were fed leaf discs treated with hypericin
exhibited a short-term, concentration-dependent
decline in enzyme activity. Longer term studies
with A. plagiata fed either the phototoxic H. perfo-
ratum, or the closely related but non-phototoxic
H. calycinum, resulted in increased CAT and GR
activity in the larvae that were fed the phototoxic
Hypericin
H 1895
plant whereas SOD activity was not significantly
different. These results suggest that CAT and GR
may be inducible defenses against phototoxins.
Similar results also have been reported for
other generalist and specialist herbivorous insects
feeding on Hypericum perforatum (Guttiferae).
The specialist insects had lower constitutive activi-
ties of two antioxidant enzymes, glutathione
S-transferase and glutathione reductase, than the
generalists. Both enzymes are considered to be
biochemical adaptations used by phytophagous
insects to attenuate the oxidative stress caused by
photosensitization.
Medicinal Use and Potential
H. perforatum is presently a source for the prepa-
ration of several herbal drugs. There is increasing
interest in use of the herbal preparation St. John’s
wort (Hypericum perforatum). Hypericin, the
major active ingredient, has many psychoactive
properties. The herbal preparation is sold as a
nutritional supplement and is recommended for
numerous conditions, including depression, anxi-
ety, insomnia and inflammation.
Hypericin, and the structurally related qui-
none, pseudohypericin, have also been described
as having potent antiviral effects, in that they could
inhibit both retroviral replication in cell culture
and could also control the symptoms of retovirus-
induced disease in mice. It has been shown to
have impressive light-mediated antiviral activities
against different viruses including human immu-
nodeficiency virus type 1 (HIV-1).
Strategies have been studied using hypericin
to inactivate enveloped viruses in red blood cell
concentrates. Photodynamically induced virus
inactivation appears promising in preventing
trans­mission of enveloped virus infections in
transfusible blood products. The results of studies
on the practical potential of hypericin as a
­photosensitizer to inactivate key enveloped viruses
in packed red blood cell concentrates (PRC) are
promising. Also, hypericin induces differentiation
1896
H Hypermetamorphosis
and apoptosis in neoplastic cells and exerts photo-
toxic effects on two cancer cell lines: (i) PC-3, a
prostatic adenocarcinoma non-responsive to andro-
gen therapy and (ii) LNCaP, a lymphonodal metas-
tasis of prostate carcinoma responsive to androgen
therapy. The results suggest that photodynamic
therapy with hypericin could be an alternative
approach to the treatment of prostatic tumors, and
could be beneficial in tumors that are non-responsive
to androgen therapy.
Effects on Domestic Animals
Recently, it has been found that the photosensitiza-
tion of grazing animals following the ingestion of
certain Hypericum species is due to the presence of
hypericin. This disease which is known as hyperi-
cism (or “bighead” in sheep) is manifested by skin
irritation and inflammation. It is most commonly
caused by the ingestion of H. perforatum.
Hypericin must be ingested by mammals to
result in hypericism. Unlike the furanocoumarins,
it does not appear to be absorbed through the outer
layer of the epidermis. After ingestion of the plants,
animals remain sensitive to sunlight for a week or
more. Skin irritation and inflammation is most
severe in regions of unpigmented skin devoid of
hair (e.g., the mouth, nose and ears). Other effects
are central nervous system disturbances and
increase in body temperature. It is, however, rarely
fatal and does not appear to deter grazing animals.
Nevertheless, sunlight-associated hyperthermia is
a consistent and rapidly developing clinical sign in
sheep intoxicated by St. John’s wort.
Mode of Action
Many secondary plant compounds are capable
of photoactivation, resulting in the production of
toxic oxygen. Early studies on mammals have
shown that the photosensitizing action of hyper-
icin requires both visible light and oxygen. Hyperi-
cin promotes Type II photodynamic reactions.
Hypericin is known to generate singlet oxygen,
and possibly other reactive species, in the presence
of light (650–700 nm). It is thought that such pho-
toproducts inactivate viruses by damaging the
membrane, since non-enveloped viruses appear to
be resistant to hypericin.
References
Aucoin RR, Philogene BJR, Arnason JT (1991) Antioxidant
enzymes as biochemical defenses against phototoxin-
induced oxidative stress in three species of herbivorous
Lepidoptera. Archi Insect Biochem Physiol 16:139–152
Colasanti A, Kisslinger A, Liuzzi R, Quarto M, Riccio P,
Roberti G, Tramontano D, Villani F (2000). Hypericin
photosensitization of tumor and metastatic cell lines
of human prostate. J Photochem Photobiol B Biol
54:103–107
Guillet G, Podeszfinski C, Regnault-Roger C, Arnason JT,
Philogene BJR (2000) Behavioral and biochemical adap-
tations of generalist and specialist herbivorous insects
feeding on Hypericum perforatum (Guttiferae). Environ
Entomol 29:135–139
Jacobson JM, Feinman L, Liebes L, Ostrow N, Koslowski V,
Tobia A, Cabana BE, Lee D, Spritzler J, Prince AM (2001)
Pharmacokinetics, safety, and antiviral effects of hyperi-
cin, a derivative of St. John’s wort plant, in patients with
chronic hepatitis C virus infection. Antimicrob Agents
Chemother 45:517–524
Samuels R, Knox P (1989) Insecticidal activity of hypericin
towards Manduca sexta larvae. J Chem Ecol
15:855–862
Hypermetamorphosis
A type of development in which there is more than
one distinct form of the larval stage. For example,
in the Meloidae, young larvae must locate a food
source, but older larvae remain and feed on that
source; thus, the long-legged, mobile young larvae
are replaced by sessile older larvae with shorter,
less functional legs.
Hyperparasitoid
An insect parasitoid that parasitizes another para-
sitoid. A secondary parasitoid.

Hyperplasia
An increase in the number of functional units of
an organ (organelles, cells, tissues), excluding
tumor formation.
Hypertely
pierre jolivet
Paris, France
The concept of hypertely, a non-Darwinian con-
cept, has been mainly developed by Lucien Cuénot
and René Jeannel in France, though originally
advanced by certain stubborn lamarckists of the
middle of the last century. It can be defined as an
excessive development of certain organs, in size
and complexity, mostly among males, and was
attributed to orthogenesis. The theory of orthogen-
esis was an alternative to Darwinian natural selec-
tion, and at one time found support in the USA
and Germany, especially among palaeontologists
and developmental biologists. Orthogenesis can be
defined as evolutionary change proceeding consis-
tently in one direction, resulting from a tendency
to accumulate similar mutations in successive gen-
erations rather than by natural selection. Thus, the
purported result of hypertely was the development
of certain organs (horns, tusks, legs, antlers, etc.) up
to the extent of a certain monstrosity.
Such a concept is very much debatable and
may be explained as the result of a normal
Darwinian selection and just part of natural bio-
diversity. Such organs are often useful or at least
harmless, and the best proof of it is that the ani-
mals endowed with them survive, multiply and do
not seem to be ill at ease with them. Beetle horns
help the males when fighting over access to females,
long legs of Sagrines help them, when mating, and
also to clutch on the stems, thoracic extensions
among the Membracidae or certain Curculionidae
can help in camouflage and, in all cases, the insects
survive well and reproduce. The extreme flatten-
ing of the Mormolyce (Carabidae) in Indonesia
Hypertely
H 1897
allow them to hide under loose pieces of bark dur-
ing the day, and similarly the aberrant tenebrionid
Cossyphus can easily slide under the bark of Aca-
cia trees in the Sudan. Hololepta plana (Histeridae)
lives under the loose bark of felled trees; it is so
flattened that its height is only a small fraction of
its width. Of course many beetles are flattened, but
this is an adaptation to their habitat. Mimicry and
camouflage are not hypertely, and may provide
protection for the animal against his predators. It
has been demonstrated recently that pronotal
horns help some beetles during ecdysis, at least
among Onthophagus (Scarabaeidae), which means
that allometry or differential growing is not just a
meaningless developmental process. An appar-
ently non-adaptive organ is often an organ not
functionally well understood, or at least an organ
that was adaptive before some change occurred in
the environment (Figs. 61 and 62).
Hypertely can be defined as “beyond the
bounds of the useful.” Some dictionaries give the
following definition: “excessive development of
certain organs, in size and complexity, among cer-
tain species, mostly among males.” The horns of
the Scarabaeidae and the mandibles of the Lucani-
dae might sometimes appear excessive, but they
help in defense, and during the fight among males
the winner is often the most well equipped with
such a horn. The thoracic prolongations of the
Membracidae can seem a bit odd, but they serve
as camouflage. The long thoracic horn of the Cra-
nopoeus (Curculionidae) mimics the seeds of the
host tree. The excessive development of certain
organs (antennae, legs) of cavernicolous (cave-
dwelling) insects compensate for the loss of eyes.
The pseudophysogastry (artificial swelling of the
abdomen, not connected with ovary develop-
ment) of some cavernicolous insects is more dif-
ficult to interpret, but it increases the isolation of
the abdomen against cold or humidity. Pseudo-
physogastry is known also among termiticolous
(living in association with termites) insects, also
living in a close and isolated world. The long ros-
trum of the Brentidae help in digging their tun-
nels. The enormous legs of certain weevils such as
1898
H Hypertely

1 2

3 4
a

c 1 2 3

2mm

e f

Hypertely, Figure 61  Some cases of hypertely (a) Hemiptera: Membracidae. 1: Bocydium; 2: C ­ yphonia;
3 and 4: Spongophorus. Tropical America; (b) Lasia nigritarsis Blanchard (Diptera: Oncididae). Male.
Tropical America; (c) Aedeagus of various Monoxia (Coleoptera: Chrysomelidae: Galerucinae). USA.
1: Monoxia puncticollis Say; 2: Monoxia debilis LeConte; 3: Monoxia sordida LeConte; (d) Dynastes
hercules (L.) (Coleoptera: Scarabeidae: Dynastinae). Male and female. Tropical America; (e) Sebethes
longipes Fabricius (Diptera: Culicidae). Male. Tropical America; (f) Acrocinus longimanus L. (Coleoptera:
­Cerambycidae). French Guyana (c, after Jolivet, 1957–59; f, after Grassé, 1949; a, b, e, after Grassé,
1951; d, after Paulian, 1935).
 Hypertely
H 1899

a b c

2mm 1mm

0
0
d f

1mm

e g

Hypertely, Figure 62  Some additional cases of hypertely (a) Diatelium sp. (Coleoptera: ­Scaphididae).
Male; (b) Rhyticephalus brevicornis Chevrolat (Coleoptera: Brentidae); (c) Nemopistha imperatrix
­Westwood (Planipennes). Equatorial Africa; (d) Diopsis tenuipes Westwood (Diptera: Diopsidae).
­Tropical Africa; (e) Phytalmia cervicornis Gerstacker (Diptera:Phytalmiidae). Male. New G ­ uinea;
(f ) Asyntona tetyroides Walker (Diptera: Platystomidae). Malaysia; (g) Sagra femorata Drury
­(Coleoptera: Chrysomelidae: Sagrinae). Thailand (figs. a and b, after Paulian, 1988; c, after Cuénot and
Tétry, 1951; d, e, f, after Grassé, 1951; g, after Jolivet and Verma, 2002).
1900
H Hypertely
neotropical Cerambycidae (Acrocinus), Malagasy
eumolpines (Arsoa; Chrysomelidae) and the big
femora of the sagrines (Chrysomelidae), are used
for holding on to the stems or mating. The sternal
horn of certain Doryphora (Chrysomelidae) is
used as a weapon in fighting between males. It
is  also evident that sensorial dimples on the abdo-
men of many beetles have their usefulness in
detection of humidity or mates. Examples are
many of those organs which appear excessively
developed, but they are not functionally
meaningless.
For Darwinists, the concept of hypertely is a
mistake of interpretation, because although the
character may look functionless, it is not without
its functional significance. The phenomenon
seems coherent and related to some function
such as sexual selection, mimicry, self defense,
clutching, lodging, etc. Some French zoologists
such as Jeannel, Cuénot, and Grassé could, in the
past, see hypertely as the result of extreme ortho-
genesis. Cases of hypertely were suggested to
include the bent tusks of the mammoths, the
bent canines of the babirussa, the teeth of the
Machairodus or Smilodon, the antlers of the Irish
deer, the horns of the Tithanotherians, the sword
of the saw fish (Pristis) and the swordfish
(Xiphias), the long spines of certain urchins
(Cidarids), the dermic armor of the Stegosauri-
ans, the complex septa of certain Ammonites,
the thoracic expansions of the Membracidae, the
horns of the Scarabeids, the mandibles of the
Lucanids, the foliaceous expansions of the legs
of certain Hemiptera or Diptera, the reduced
hind wings of the Nemopteridae, and even the
long legs of the Tipulidae. However, these seem-
ingly unusual structures can be explained. For
example, the long legs of those Diptera prevent
them from getting captured on spider webs when
they dance rhythmically, and the mandibles of
the Lucanidae and the horns of the Scarabeidae
help them when fighting (and those organs have
some other functions too, recently decrypted).
Some beetles have a very elongated body, but
this form is useful for digging and living in
t­ unnels. Others, as the Cychrus (Carabidae) have
the head and pronotum elongated and narrow,
but this is an adaptation used to penetrate snail
shells. The long proboscis of certain Diptera or
moths are needed to penetrate some deep flow-
ers. All those modifications of the morphology
seem to be ­functional specializations, not mean-
ingless exaggerations.
All modern authors see in the so-called hyper-
tely of the carabeid horns a case of allometry,
sometimes under the influence of hormones or
food. Individuals are able to express different mor-
phologies in response to environmental condi-
tions during their development. Recently, it was
suggested that pronotal horns among Onthopha-
gus are crucial during ecdysis of the larval head
capsule during the larval-to-pupal molt. This func-
tion in molting seem unrelated with fighting and
defense, but all criteria are positive and useful for
the insect.
Additional examples of exaggeration include
organs used in reproduction; often, the male geni-
talia of certain beetles are very big, while other
species of the same genus have a normal aedeagus.
That is the case among the galerucines of the genus
Monoxia (Chrysomelidae), where only one spe-
cies, Monoxia puncticollis, has an enormous intro-
mittent organ, but its body size is not much bigger
than the body of other species with a normal aede-
agus. The mating is done and successful with the
female of M. puncticollis, which probably has some
kind of adaptation. There is also the case of Hydro-
chus interruptus (Hydrophylidae), an aquatic bee-
tle, which is provided with so complex an aedeagus
that some entomologists have doubted its real sex-
ual function. Very often, the beetles or other insects
are gifted with so complicated aedeagi that one
wonders how they work. But such aedeagi func-
tion, and the insect reproduces normally. Also,
sometimes the spermatozoa are enormous. This is
the case of many Alticinae (Chrysomelidae) or the
Drosophila flies (Drosophilidae), for instance.
However, it appears that, in the case of the Altici-
nae, the role of the long sperm is in forming a vag-
inal plug. It seems also that the males of Drosophila
 Hypodermosis in Deer
H 1901

which have developed those long spermatozoa Hypertrehalosemic Hormone

have an increased chance to fertilize the eggs (Hth)
successfully.
As far as we can see, the concept of hypertely A peptide hormone important in the regulation
is obsolete. All those so-called exaggerated of carbohydrate metabolism.
­morphologies have a function and the animals
survive, multiply and they do not seem to suffer
from those “monstrosities.” As a rule, handicapped Hypertrophy
specimens are eliminated and only the fit ones
survive. A non-adaptative organ is an organ not An increase in the size of the functional units of
understood. So-called hypertelic insects are just an organ (organelles, cells, tissues), excluding
part of the biodiversity, having evolved through tumor formation.
selection.

Hypha (pl., hyphae)

References A strand of the threadlike mycelial tissue of fungi.

Carayon J (1977) Insémination extra-génitale traumatique.

Traité de Zoologie, Masson publ., Paris 8
(5A):351–390
Hypochthonellidae
Cuénot L, Tétry A (1951) L’Evolution Biologique. Masson,
Paris, 592 pp A family of bugs (order Hemiptera, suborder Ful-
Eberhard DJ (1985) Sexual selection and animal genitalia. goromorpha). All members of the suborder are
Harvard University Press, Cambridge, MA, 244 pp
Emlen DJ, Nijhout HF (2000) The development and evolution
referred to as planthoppers.
of exaggerated morphologies in insects. Ann Rev Entomol
45:661–708
Gould SJ (1977) Ever since Darwin. Reflexions in Natural Hypodermosis in Deer
History. Penguin Books, Harmondsworth, England,
285 pp
Javier martínez, ignacio navarrete,
Grassé PP (1949) (1951) Traité de Zoologie. Anatomie, Systé-
matique, Biologie. Insectes. IX & X (1 & 1). Masson, david reina, santigo hernández
Paris, 1117 + 1949 pp Universities of Cordoba and Extremadura,
Jolivet P (2004) Inverted copulation. In: Capinera J (ed) ­Madrid, Spain
Encyclopedia of entomology, vol 2. Kluwer Academic
­Publishers, Dordrecht, The Netherlands, pp 1208–1212
Moczek AP, Cruickshank TE, Shelby A (2006) When ontog- Hypodermosis is a common myiasis produced by
eny reveals what phylogeny hides: gain and loss of horns flies of the genus Hypoderma (Diptera: Oestri-
during development and evolution of horned beetles. dae), affecting a wide variety of hosts, but spe-
Evolution 60:2329–2341
Moczek AP, Emlen DJ (2000) Male horn dimorphism in the cially important in domestic animals like cattle
scarab beetle, Onthophagus taurus: do alternative repro- and goats. It is spread over almost all the north-
ductive tactics favour alternative phenotypes? Anim ern hemisphere, and these flies produce serious
Behav 59:459–466
economic losses in hosts, especially in herds
Paulian R (1935) Le polymorphisme des mâles de Coléoptères.
Hermann, Paris, 35 pp reared for meat and dairy production. Little
Virkki N, Bruck T (1994) Unusually large sperm cells in research has been directed, however, at wild and
­Alticinae: their formation and transportation in male semi-wild animals such as deer. Nonetheless,
genitalia system and their evolution. In: Jolivet P, Cox ML,
Petitpierre E (eds) Novel aspects of the biology of
existing studies have pointed out just how seri-
Chrysomelidae, Kluwer Academic Publishers, ­Dordrecht, ous this infestation may be in these hosts, because
The Netherlands, pp 371–381 widespread parasitosis currently affects deer in
1902
H Hypodermosis in Deer
numerous Palaearctic countries. The situation is
truly worrying, due to the high degree of preva-
lence and intensity of parasite infestation.
Effects of Hypodermosis
Little is known about the infestation of deer,
and few studies have discussed the influence of
environmental factors on the biology of flies and
their resulting distribution. At the same time, the
pathogenicity of larvae for host deer remains
unknown. Although it is patently clear that, in
deer, this disease should be classed as critical,
since it affects:
Deer Health
The Hypoderma life cycle in deer is still not well
known, but due to the similarities with cattle myi-
asis, it is thought to have an internal migratory
cycle that affects the health of the host. There is a
decrease in meat production, retarded growth and
reduction in skin quality (Fig. 63).
Hypodermosis in Deer, Figure 63  Myiasis in deer caused by Hypoderma.
Rural Development in Poor Areas
Deer hypodermosis affects the economic develop-
ment of many poor areas due to repercussions on
the hunting-based economy. In social terms, hunt-
ing is a developing force in depressed areas, where
it provides a major source of income through the
employment of local labor.
Deer Farming
This is a growing farming activity, and venison
offers an alternative to beef, pork and lamb.
Species and Hosts of Hypoderma
spp.
Several Hypoderma species are found in deer, but
the three main parasites are H. diana, H. actaeon
and H. tarandi. H. diana is euryxenic and affects,
in addition to red deer (Cervus elaphus), a range of
other hosts including roe deer (Capreolus capreo-
lus), fallow deer (Dama dama), elk (Alces alces)

and reindeer (Rangifer rangifer), and lives in a
great diversity of ecosystems.
Host specificity is strong in H. actaeon and
H. capreolus, as the former only affects red deer
and the latter roe deer.
Morphological differentiation between H. diana
and H. actaeon is made according to the pattern of spine
morphology and morphology of spiracular plates.
H. (Oedemagena) tarandi shows greater
­specificity, affecting only R. rangifer, and is found
within the subarctic region.
Biology
Biology of deer-infesting Hypoderma spp. is still a
relatively unknown aspect of myiasis. Some par-
ticulars are important in deer hypodermosis. For
example, H. diana is present in a great variety of
habitats, and also of hosts, and this departs from
the principle of host specificity of warble flies,
maintained by a complicated equilibrium with its
host by means of internal regulatory ­systems. It is
spread throughout Europe and Asia, from 30° to
60° north, and lives in several different ecological
zones such as mixed, deciduous and coniferous
forests, wooded steppes and wetlands, overlapping
the territory of its hosts and adapted to the wide
variety of ecosystems in which live the parasite’s
hosts.
Flight and oviposition of female flies are con-
ditioned by air temperature and ambient light, so
they are most active at midday. In contrast, their
endogenous cycle has yet to be fully described,
although some authors claim that they arrive at
the dorsal area via the spinal canal, exhibiting a
cycle similar to that of H. bovis. Resistance and
susceptibility to infection has to be studied, as
previous studies have demonstrated that the
extent of ­parasitism and prevalence are higher in
younger animals, possibly due to a measure of
resistance built up through repeated contact with
the parasite.
Pupation is highly influenced by the nature of
the land, pupae being more viable when produced
Hypodermosis in Deer
H 1903
in areas where there is at least some grass. This
viability decreases depending on to what extent
the land freezes or frosts over.
Hypoderma tarandi is more specific and is
found in just one host (R. rangifer); it has thus
adapted itself to live in the same habitat as its host,
usually subarctic regions. Its biology has been
studied in great detail due to the importance of the
reindeer in cold northern countries. Two different
habitat are occupied by these flies, one of wooded
regions, and other of wide open or mountainous
areas with plentiful water courses. Usually only the
female approaches the host; the male rarely does
so, and is usually found in sunny, rocky areas. The
flight range of females is absolutely astonishing, as
they can fly as far as 600–900 km. Males, however,
tend to fly for shorter distances and times, although
they may achieve a radius of 400 km.
The degree of parasitism in male deer is usu-
ally higher than in females and castrated animals;
thus, resistance to this disease is greater in females
than in males. Prevalence is higher in herds which
did not move to new pastures after calving than in
those continually changing their grazing grounds.
When deer are led to new pastures, the number of
infested animals and the degree of parasitism
drops substantially.
Hypoderma actaeon is confined to one host,
red deer (Cervus elaphus), and it seems to be
restricted to Europe. Little is known about the
biology of this parasite, but it has been observed
that yearling hosts are more parasitized than older
ones, and it suggests the development of resistance
by repeated contact with the parasite. The internal
life cycle of H. actaeon lasts up to 9 months, as
subcutaneous stages are found from September to
May, with the presence of first instar larvae for a
period of 2–3 months, and then the second (L2)
and third (L3) instars.
Chronobiology
The seasonal dynamics of H. diana adapted to
central European countries are as follows: larva
1904
H Hypodermosis in Deer
migration and growth take place in the host until
April, pupation lasts for 26–33 days, and flying is
observed in May and June. In eastern countries
(Czech Republic), L1 larvae develop from November
to March, L2 from December to April, and L3
from January to June. Flies have been observed
from April to June. By contrast, in southern European
countries such as Spain, this cycle is brought
forward by 2–2.5 months. L1 may be observed
September to December, L2 October to January,
and L3 onwards of December.
Hypoderma tarandi, owing to its northern
ecology, does not develop until later, beginning in
October and with L3 appearing June to July; flying
takes place between June and September.
Chronobiology of H. actaeon is similar to
that described to H. diana, with L1 larvae devel-
oping in September to November, second instar
larvae from September to December and L3 from
December to May.
Prevalence
Hypoderma diana is widely distributed through-
out Europe, and cases of parasite infestation as
high as 100% have been reported, such as those
recorded in Russia. In Bulgaria, parasitism ranges
from 48 to 66%, whereas in Spain there is a mean
of 90%.
The other important species in red deer is H.
actaeon, which reach very high levels of prevalence
in southern and central Spain, ranging from 44 to
92%, with a mean intensity of parasitization of
near 40 larvae per animal (1–317).
The extent of parasitism with regard to the
number of larvae per animal varies greatly in H.
diana and H. actaeon. In Spain, values ranging
from 1 to 400 larvae per animal have been recorded,
where over half of the animals contained more
than 50 larvae. In the former Czechoslovakia, cases
have been reported of up to 300 larvae per animal,
in Germany 300–500 larvae, in Bulgaria up to 150
larvae, and in France up to 200. H. tarandi has also
reached levels of 99.9%.
Other Negative Effects of
Hypodermosis
It has been shown that H. diana can infest other
hosts in addition to deer. In the United Kingdom,
cases have been reported of sheep living near
deer-populated forests being affected. Cases of
ophthalmomyiasis produced by H. tarandi have
been described in children, in dogs and in horses,
and always in the vicinity of deer-populated
areas.
The cost of the damage caused by Hypoderma
spp. is difficult to calculate although, as occurs in
bovines, losses are incurred both in reduced meat
yield and in depreciation of skins and horns/
antlers.
Control
It is never easy to develop strategies for the control
of disease in wild and semi-wild animals, and deer
are no exception. Any such controls also must be
based on a total respect for the environment, since
the quality of their surroundings plays an impor-
tant role in the welfare of not only deer but all wild
animals living on the same biotope.
It should be borne in mind that ecological
or “organic” meat such as venison, supplied to a
gourmet market, must satisfy sophisticated orga-
noleptic criteria and its quality must never suffer.
Thus, the use of medicaments and pharmacological
substances has to be strictly controlled in order
not to affect meat quality.
There are only a few studies about the treat-
ment of warble flies in deer, but we can assume
that the same drugs used in other species should
be useful in deer, i.e., some organophosporous,
avermectines and milbemicynes.
Handling practices may be modified through
a knowledge of the parasite cycle and ecology.
Animals could enjoy greater protection by being
herded into corrals, or at least into the shelter of
shaded areas to reduce the risk of infestation when
adult flies are active.
 Hypostome
H 1905

In domesticated animals or semi-domesti- San Miguel JM, Álvarez G, Luzón M (2001) Hypodermosis of
red deer in Spain. J Wildlife Dis 37:342–346
cated animals living on the range, such as reindeer,
Zumpt F (1965) Myasis in man and animals of the old world.
the rate of infestation may be reduced when, after Butterworths, London, UK
giving birth, the herd migrates to fresh pastures or
is led to new pastures away from those frequented
during the muddy season. The aim is to distance Hypogaeic
the deer from the pupae and flies. These measures
are not ­particularly effective, however, so greater Living primarily underground, or at least beneath
control of parasitism will still depend on pharma- a layer of material on the soil surface. (contrast
cological treatment. with epigaeic)
For free-range deer, and only when they
­cannot be captured for the administration of treat-
ment, food may be supplemented with oral forms
Hypogastruridae
of antiparasite preparations. The problem arises,
A family of springtails (order Collembola). They
however, in choosing a suitable vehicle for the
commonly are known as elongate-bodied
anti-parasitic treatment, because it cannot be
springtails.
freely distributed around the environment due to
 Springtails
the danger of its being consumed by other ani-
mals. Neither could the ingested volume be con-
trolled. Experimental trials with rafoxanide and Hypognathous
ivermectin in the daily salt pellets required by the
animals’ diet have yielded promising results. An insect that has the head pointing forward and
Farm-raised animals are obviously easier the mouth pointing downward. It is also known as
to control, and a program of preventative medi- orthognathous. This is considered to be the ances-
cine may be applied, with regular treatment in tral condition for insects.
accordance with the local chronobiology of  Mouthparts of Arthropods
Hypoderma.
 Flies
 Veterinary Pests and Their Management Hypopharynx
A tongue-like structure in the buccal cavity on the
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Hernández-Rodríguez S, De La Fuente López, C,
Hypostomal Bridge
Alunda JM, Hall MJR (1998) Comparative scanning
electron microscopy of third instar Hypoderma spp. The portion of the head capsule that closes off the
Med Vet Entomol 12:181–186 ventral area of the foramen magnum, near the
Martinez-Moreno FJ, Navarrete I, Hernandez-Rodriguez S
base of the labrum.
(1995) Deer hypodermosis in the south of Spain. In:
Tarry DW, Pithan K, Webster K (eds) Improvements in  Head of Hexapods
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E.C., Luxembourg, Luxembourg, pp 115–124
Minar J (1982) A contribution to the knowledge of distribu- Hypostome
tion and ecology of botflies (Oestridae) and warble flies
(Hypodermatidae) in the cervids. Folia Facultatis
Scientiarum Naturalium Universitatis Purkynianae In ticks, this is a denticle-bearing ventral structure
Brunensis, Biologia 23:87–92 occurring parallel to, and between, the palpi.
1906
H Hypsypterygidae

Hypsypterygidae Hystrichopsyllidae

A family of bugs (order Hemiptera). A family of fleas (order Siphonaptera). They some-
 Bugs times are known as rodent fleas.
 Fleas