to Arthropod Bites
John H. Klotz, Jacob L. Pinnas, Stephen A. Klotz, and Justin O. Schmidt
Two French scientists, Charles Richet and Paul Portier, codiscovered
anaphylaxis in 1901. The lion’s share of the
credit went to Richet, an eminent physician and Professor of
Physiology at the University of Paris, while Portier was an assistant
in the Laboratory of Physiology at the Sorbonne. A history of their
discovery given from the perspective of Portier’s contributions is
well covered by May (1985).
Their collaboration began as guests on the oceanographic research
vessel of Albert I, Prince of Monaco, as commemorated by a
stamp (Fig. 1). The prince and his scientific director suggested the
topic for their research on this scientifically historic cruise. They were
to conduct experiments on the toxicity of venoms from the Portugese
man-o-war, Physalia physalis.
Figure 1. Postage stamp issued by Monaco in 1953 to commemorate
the discovery of anaphylaxis. This figure was published in the Journal of
Allergy and Clinical Immunology, Vol. 110, Cohen, S.G. and M. Zelaya-
Quesada: Portier, Richet, and the discovery of anaphylaxis: a centennial,
p. 333, Copyright Elsevier (2002).
In the definitive experiment conducted after they returned to
Paris, Richet and Portier exposed two dogs to weak doses of sea
anemone actinotoxin and then repeated the injection at various time
intervals. No reaction was noted until an injection 26 days after the
beginning of the experiment, when both dogs became extremely
ill and died shortly thereafter. Richet (1913) proposed two factors
that were necessary and sufficient to cause an anaphylactic reaction:
“increased sensitivity to a poison after previous injection of the same
poison, and an incubation period necessary for this state of increased
sensitivity to develop.”
To name this reaction, Richet first proposed the term “aphylaxis,”
later coining the term anaphylaxis (“without protection”) because it
was more euphonious. For their studies on hypersensitivity reactions,
Richet was awarded the 1913 Nobel Prize in Medicine or Physiology,
while Portier, although his contribution was significant, did not
share in the Prize and was barely mentioned in the Nobel address.
Given the academic tradition at that time, it was not unusual for the
more distinguished senior scientist to overshadow the lesser-known
junior scientist, and Portier, a humble man, apparently did not feel
slighted and remained friendly with Richet until the latter’s death
in 1935. Portier made other important scientific contributions, including
entomological studies concerning the physiology of aquatic
insects and a treatise on the biology of butterflies published in 1949,
when he was 83.
Anaphylaxis described. Current definitions of anaphylaxis
reflect advances in our understanding of its physiological basis: an
acute systemic allergic reaction resulting from the release of chemical
mediators following an immunologic reaction that is typically
mediated by immunoglobulin E (IgE). IgE is one class of antibodies
produced by the immune system in response to foreign substances.
Individuals who suffer allergic reactions produce greater quantities
134 American Entomologist • Fall 2009
of IgE toward allergens to which they have been sensitized and have
poorer regulation of responses to those allergens.
A sensitization phase in which the individual is “set up” for the
reaction (Frazier and Brown 1980) directs lymphocytes to recognize
invading allergens as foreign and activates them to produce IgEantibodies.
The lymphocytes then release these antibodies, which
subsequently bind to receptor sites on mast cells and circulating
basophils. The mast cells contain histamine and are located in body
tissues such as the respiratory and gastrointestinal tracts, the heart,
and the mucous glands and skin.
In the sensitized individual, re-exposure to the offending allergen
sets in motion a cascade of biochemical events. The allergen crosslinks
neighboring IgE antibodies on mast cell and basophil surfaces,
which alters the cell membrane and leads to the release of histamine
and other chemical mediators. These mediators are capable of
contracting smooth muscle in the airways and intestines, as well as
dilating blood vessels and increasing vascular permeability.
These pathophysiological events may manifest as hives or skin
swellings (angiodema), labored breathing, dizziness, or shock.
The time interval between a bite or sting and appearance of these
symptoms is often short, usually only several minutes. Reactions to
stings by Hymenoptera, for example, vary within a continuum ranging
from minor to severe. In most cases the reaction is limited to a
welt (hive)—a reddened, tender area that causes burning and pain
for an hour or two (Greene 2005). In a large local reaction, there is
pronounced swelling; sometimes an entire leg or arm will swell from
a sting on the toe or finger.
The most common systemic reactions are cutaneous: the individual
breaks out in hives or urticaria. There is the rare fatal anaphylactic
reaction, in which the individual typically dies within 30
to 60 minutes after the sting due to respiratory and cardiovascular
complications. These are not toxic reactions, but allergic reactions
to proteins in the venom (toxic reactions take hours to days, unless
several thousands of stings are received).
Causative agents of Anaphylaxis. Since its discovery in the early
1900s, other causative agents of anaphylaxis in addition to venom
have been implicated, including foods, which are the most common
cause of anaphylaxis outside of the hospital setting (50-100 deaths/
year), medications (especially antibiotics), latex, vaccines, hormones,
and sometimes even exercise when associated with a particular food
(Kemp 2001; Fireman 1999).
Arthropods are by far the most common cause of anaphylaxis
due to animal bites or stings, and the insects (particularly Hymenoptera)
make up the majority of these cases (see Table 1 for cases of
anaphylaxis caused by arthropods that have been determined to be
IgE-mediated and Table 2 for other reported cases of anaphylacticlike
reactions, so called because these reports lack the definitive in
vitro or in vivo tests to demonstrate IgE-mediation).
Hymenoptera. It is speculated that the earliest case of an allergic
reaction dates back to ancient Egypt (3300 to 2640 BCE), when the
Pharaoh Menes supposedly suffered a fatal anaphylactic reaction
when stung by a “kheb” during a journey to the “Western Isles”—possibly
referring to Britain (Harper 1980). In ancient Egyptian, kheb
means hornet or hippopotamus, and to the dismay of the allergists
who favor this historical footnote as to the origin of their profession,
some Egyptologists have implicated the hippo in the death of King
Menes, especially given its abundance in the Nile (Harper 1980).
Table 1. Anaphylactic reactions to arthropod bites and stings (evidence
for IgE-mediation) (Klotz et al. 2008)
Scientific Nomenclature (Common names in parentheses were reported
to induce the reaction)
Genus: Vespula (ground-nesting yellow jackets)
Dolichovespula (aerial-nesting yellow jackets)
Polistes (paper wasps)
Genus: Apis (honey bees)
Bombus (bumble bees)
Genus: Solenopsis (fire ants)
Pogonomyrmex (harvester ants)
Myrmecia (bulldog ants)
Pachycondyla (Asian needle and Samsum ants)
Formica (wood ants)
Genus: Triatoma (kissing bugs)
Genus: Chrysops (deer flies)
Tabanus (horse flies)
Family: Simuliidae (black flies)
Hippoboscidae (louse flies)
Genus: Glossina (tsetse flies)
Genus: Thaumetopoea (pine processionary caterpillars)
Genus: Ixodes holocyclus (Australian paralysis ticks)
Ixodes pacificus (western black-legged ticks)
Genus: Argas (pigeon ticks)
Genus: Centruroides (bark and common striped scorpions)
Class: Chilopoda (centipedes)
About 1% of children and 3% of adults are allergic to stings of
Hymenoptera, and at least 40 fatalities occur each year in the United
States, with many victims having no previous reactions to stings
(Golden 2003). Sting allergies in the U.S. are most commonly due to
yellow jackets (Vespula and Dolichovespula) and honey bees (Apis),
followed by fire ants (Solenopsis) and paper wasps (Polistes), and less
frequently, harvester ants (Pogonomyrmex), hornets (Vespa), bumblebees
(Bombus), and sweat bees (Halictidae) (Schmidt 1992).
All of these insects, except some sweat bees, are social and most
are characterized by central place foraging: worker bees, wasps, and
ants leave a nest to find food and then return with it to provide for
the colony. The distance traveled to obtain food varies from a few
meters for ants to several kilometers for bees. The daily range of
the colony’s forays defines its home range or territory. As the colony
grows in size, its territory expands to accommodate the increasing
American Entomologist • Volume 55, Number 3 135
Table 2. Anaphylactic-like reactions to bites and stings of arthropods
(lacking laboratory evidence for IgE-mediation) (Klotz et al. 2008)
Scientific Nomenclature (Common names in parentheses were reported
to induce the reaction)
Family: Halictidae (sweat bees)
Genus: Pseudomyrmex (twig ants)
Rhytidoponera (green-head ants)
Genus: Cimex (bed bugs)
Genus: Culicoides (punkies)
Genus: Symphoromyia (snipe flies)
Genus: Thereva (stiletto flies)
Genus: Chelepteryx (white-stemmed gum moths)
Genus: Megalopyge (puss caterpillars)
numbers of individuals. Their territorial defense is maximal at the
nest, which they tenaciously defend against any intruder.
In the aculeate Hymenoptera (wasps, bees, and ants), the ovipositor
of females has been modified into a stinger, an adaptation that
enables these insects to defend their nest against potential predators
that would exploit such a concentrated source of food (Schmidt
1986). In many species of ants the stinger is absent or vestigial, but
they still possess potent defensive secretions.
Although ants are not generally appreciated as causes of anaphylaxis,
there are a growing number of species in the U.S. that have been
reported to cause this medical emergency (Table 3). Most notorious
is the red imported fire ant, Solenopsis invicta, which is widespread
in the southeastern U.S. and continuing to expand its range. In some
infested urban areas > 50% of the population is stung per year (de
Shazo et al. 1990) and as much as 17% of the population is sensitized
(Caplan et al. 2003). At least 80 deaths have been attributed
to imported fire ants (Rhoades et al. 1989). Particularly vulnerable
are the elderly and infirm in nursing homes or hospitals, unable to
defend themselves from attack because of limited mobility.
Infants represent another vulnerable group to fire ant stings.
Two fatalities attributed to southern fire ants, Solenopsis xyloni, were
babies less than a year old (Coarsey 1952; Klotz et al. 2004). Nonfatal
systemic reactions have been reported for two other native fire ant
species, S. aurea and S. geminata (Hoffman 1997), although the native
fire ants are generally less aggressive than imported fire ants.
Fire ant stings are characterized by an intense burning sensation
due to alkaloid compounds (piperidines) in the venom. Each species
has its own unique blend of these compounds, but imported fire
ant stings are the most severe and typically cause the formation of
Harvester ants possess the most toxic of all insect venoms; in
fact, drop for drop, it is more toxic than rattlesnake venom (Schmidt
Table 3. Ant species and their geographic distribution in the continental
U.S. that have been reported to cause anaphylactic or anaphylactic-like
reactions (Klotz et al. 2005b)
Scientific Name Range
Solenopsis invicta Southern US, New Mexico, California
Solenopsis xyloni Southern and southwestern US
Solenopsis aurea Southwestern US
Solenopsis geminata Southern US
Pogonomyrmex rugosus Western Texas and Oklahoma into
Pogonomyrmex maricopa West Texas into southern California
Pogonomyrmex barbatus Kansas south to Texas into Arizona
Pseudomyrmex ejectus Southern US
Hypoponera punctatissima Florida, Northeast, Pacific Northwest
Pachycondyla chinensis Georgia to Virginia
1978). The sting has been described as “ripping muscles or tendons”
and “turning a screw in the flesh” (Schmidt 1986). A neurotoxic component
in the venom causes gooseflesh and sweating to occur at the
sting site, and often victims develop pain and tenderness in nearby
lymph nodes. Allergic reactions to the sting have been reported.
For example, of eight patients treated for stings over a one-year
period in Tucson, Arizona, four had large local reactions, and four
were treated for anaphylaxis (Pinnas et al. 1977). The two species
responsible, the Maricopa harvester ant, Pogonomyrmex maricopa,
and the rough harvester ant, P. rugosus, are commonly found in urban
areas of Tucson (Klotz et al. 2005a). Schmidt et al. (1984) determined
that there was significant cross-sensitivity of patients to the various
species of harvester ants.
Isolated cases of anaphylactic and anaphylactic-like reactions
have been reported in the U.S. for ants belonging to several genera.
One case reported from South Carolina resulted from the sting of an
unidentified species of Tetramorium (Majeski et al. 1974). Two cases
in the southeast were due to stings by Pseudomyrmex ejectus (Klotz
et al. 2005b), a twig-inhabiting ant that lives in small colonies with
This species does not have a sting; nevertheless, it caused an anaphylactic
reaction through its bite (Schmid-Grendelmeier 1997). In addition
to biting, ants in this genus spray formic acid that is produced
in the venom gland along with other compounds.
Hemiptera. In comparison to the stinging Hymenoptera, biting
insects cause far fewer allergic reactions. By far the most frequent in
the U.S. are allergic reactions to bites from kissing bugs (Triatoma and
Paratriatoma spp.). For example, in one small community in Santa
Barbara County, California, about 7% of the population has been
sensitized to bites of the western conenose bug, Triatoma protracta
(Marshall et al. 1986). Another species, T. rubida, is more common
in central and southern Arizona and is particularly a problem in the
foothills of Tucson, where most of the allergic reactions are reported
for this species (Pinnas et al. 1986). Interestingly, there is little to
no antigenic cross-reactivity between these two species (Pinnas et
There are 14 other species of Triatoma that are found throughout
the lower two-thirds of the U.S. (Vetter 2001). All are blood-suckers
that normally parasitize wood rats, opossums, raccoons, and armadillos
(Vetter 2001). During the spring and early summer dispersal,
T. rubida and T. protracta sometimes enter homes and feed on the
occupants as in the following case (Klotz et al. 2006):
A 45-year old woman had four severe reactions to an insect bite.
She found the insect in bed each time and from the description, it fit
the picture of a kissing bug. She never felt the bite, but noticed her
heart rate increasing and felt hot. One of the authors (JP) attributes
the rapid heartbeat to the adrenal gland’s response to elevated histamine
levels, which sometimes can “self-treat” and thereby result in
non-life-threatening reactions. In two instances, she lost consciousness
and during one episode, she had a seizure. Her son captured a
specimen from her bed that was identified as T. rubida.
Although bedbugs are an emerging problem in the U.S., anaphylactic
reactions to their bites, are rare (Parsons 1955).
Diptera. Blood-sucking flies, including horse flies (Tabanus), deer
flies (Chrysops) (Freye and Litwin 1996; Hemmer et al. 1998; Hrabak
and Dice 2003; Wilbur and Evans 1975), black flies (Simuliidae)
(Hoffman 1987), tsetse flies (Glossina) (Stevens et al. 1996), louse
flies (Hippoboscidae) (Vidal et al. 2007), and mosquitoes (Culicidae)
(McCormack et al. 1995), have caused anaphylactic reactions.
Considering the number of people bitten by mosquitoes, there are
surprisingly few reports of anaphylactic reactions, but large local
reactions are not uncommon (Engler 2001).
Anaphylactic-like reactions to punkies (Culicoides) (Hoffman
1987), snipe flies (Symphoromyia) (Turner 1979), and stiletto flies
(Therevidae) (Smith 1979) have been reported. The larvae of punkies
are aquatic or semiaquatic and the adults do not travel far from
where the larvae live (Triplehorn and Johnson 2005). Most snipe
flies do not bite, but several species of Symphoromyia do bite and
are common in western mountain and coastal regions of the U.S.
(Triplehorn and Johnson 2005). In the case of the stiletto fly, it was
the larva that bit the victim. The adult flies are uncommon but the
predaceous larvae can be found in sand or decaying wood (Triplehorn
and Johnson 2005).
Lepidoptera. There are several families of Lepidoptera with caterpillars
possessing stinging hairs, some with venoms that can cause
anaphylactic reactions in susceptible individuals. These include the
pine processionary caterpillar, Thaumetopoea pityocampa, so called
because of the long lines they form, sometimes with hundreds of individuals
following a lead caterpillar to and from the nest (Sbordoni
and Forestiero 1985). Their hairs, which can be airborne, penetrate
the skin and release a toxic substance (Vega et al. 1999, 2000), or may
be inhaled or ingested. The most common reaction is a cutaneous
lesion; however, in northwestern Spain, as many as 40% of patients
diagnosed with occupational urticaria suffered anaphylactic reactions
(Vega et al. 2004). There are also reports of systemic reactions
in children (Shkalim et al. 2008).
Anthelid larvae and frequently their cocoons are protected by
stinging hairs. There are about 100 species native to Australia and
New Guinea, and the larger members belong to the genus Chelepteryx
(Sbordoni and Forestiero 1985). The caterpillar of the whitestemmed
gum moth (Chelepteryx collesi) incorporates urticating
hairs into its cocoon. The hairs, which point out, readily penetrate
human skin and have caused anaphylactic-like reactions (Mulvaney
et al. 1998).
In Texas from 1955 to 1959, there were 54 reported cases of
stings by caterpillars: 47 were localized reactions, and the others
anaphylactic-like reactions (Micks 1960). The caterpillar was identified
in 43 of these cases and determined to be Megalopyge opercularis.
Commonly known as puss caterpillars, they are covered with soft
brown hairs with poison spines beneath that can penetrate the skin,
causing severe reactions (Borror et al. 1976).
Acari. There are several reports of anaphylactic reactions to bites
from hard ticks (Ixodidae), most notably the Australian paralysis tick,
Ixodes holocyclus (Gauci et al. 1989). Ticks caused approximately
0.7% of the reported allergic reactions to arthropod stings and bites
in Queensland, Australia (Solley 1990). In other parts of the world,
there are fewer reports: a 73-year old man who suffered recurrent
anaphylaxis due to bites from Ixodes pacificus (Van Wye et al. 1991a,
b), a widely distributed species in western North America; and cases
in western Europe involving Ixodes ricinus (Moneret-Vautrin et al.
1998) and Rhiphicephalus spp. (Acero et al. 2003; Valls et al. 2007).
The pigeon tick, Argus reflexus, is a much more common cause
of anaphylactic reactions in Europe (Hilger et al. 2005; Rolla et al.
2004). It is a soft tick (Argasidae) that is a temporary parasite of wild
and domesticated pigeons. When its normal host is absent, the ticks
may migrate into households and bite the human occupants, some
of whom may be allergic.
Scorpions. Fatalities due to scorpion stings are by some estimates
as high as 50,000 deaths per year worldwide (White 1995). These
are mostly due to toxic reactions. Even in certain parts of the U.S.,
scorpion stings are common. For example, in Arizona (excluding the
greater Phoenix area), there were 4,655 scorpion stings reported
over a two-year period from 2002-2004 (Klotz et al. 2005a).
Fatalities due to scorpion stings are rare in the U.S., but five deaths
were recorded from 1950-1954, one of which was an anaphylactic-like
reaction (Parrish 1959). In 2001 in Arizona, a woman died
from an anaphylactic-like reaction to a sting by the bark scorpion, C.
exilacauda (Boyer et al. 2001). It is the deadliest species in the U.S.
and is mainly found in Arizona (Curry et al. 1984). IgE-mediated
anaphylaxis to its sting has also been reported (Chase et al. 2002).
Although anaphylactic reactions to their stings are rare, the venom
of the common striped scorpion, Centruroides vittatus, is reportedly
cross-reactive with imported fire ants, S. invicta (Nugent et al. 2004).
The geographic distribution of striped scorpions and imported fire
ants overlap, possibly placing many more people at risk for allergic
American Entomologist • Volume 55, Number 3 137
Chilopoda. Centipedes have a pair of poison claws on the first
segment behind the head that can inflict painful bites. Skin prick tests
with centipede venom were positive in three patients with systemic
allergic reactions to their bite (Harada et al. 2005).
Treatment of Anaphylaxis
Physicians treating allergies can offer their patients three options:
(1) medications, (2) immunotherapy, and (3) avoidance of the
allergen (Fireman 1999).
(1) Medications. Given the speed of an allergic reaction to a bite
or sting, immediate medical attention is critical. Epinephrine and
antihistamines may be life-saving when administered early during
an anaphylactic reaction. A prescription from a physician for selfinjected
epinephrine is advisable for sensitized individuals who are
at risk of life-threatening reactions. They should carry and know how
to administer a preloaded syringe containing two doses; the second
dose may be needed in some severe reactions.
The antihistamines act by binding to the receptor sites on target
cells, thereby blocking the effects of histamine. Epinephrine has
multiple anti-inflammatory effects. Methylprednisolone, a corticosteroid
with broad anti-inflammatory properties, is often administered
and is long-acting, but it requires hours before reaching maximum
effectiveness. Its short-term benefit is questionable.
(2) Immunotherapy. For the more common causes of insect sting
allergy (yellowjackets, honey bees, and imported fire ants), immunotherapy
is available and involves repeated injections of increasing
doses of the venom extract, or in the case of imported fire ants,
whole body extract. Possible mechanisms for the beneficial effects
of immunotherapy include activation of lymphocytes to produce IgG
blocking antibodies, which have a high affinity for the allergen and
can prevent it from binding to mast cells, and production of suppressor
T lymphocytes, which suppress IgE production of B lymphocytes.
Unfortunately, for less common causes of allergic reactions to bites
and stings, commercial extracts for immunotherapy are not available.
Nevertheless, some allergists have developed immunotherapy for
these so-called “orphan insects.” These are not generally available
and involve only a small segment of the population—usually only
their own patients.
(3) Avoidance of allergen. Correct identification of the offending
arthropod is critical for understanding its biology, which may provide
useful information in preventing future accidental contacts. It is often
helpful to the healthcare provider to contact an entomologist or pest
management professional for help in identification and elimination
or avoidance of the offending arthropod.
With ever-increasing urban development and sprawl into natural
habitats, there is a growing problem of bites and stings by a variety
of arthropods with the potential to induce allergic reactions. It is
important that the causative agents be identified and reported to
state or local poison control centers so that a record is maintained.
Physicians, entomologists, pest management professionals, and the
general public need to be made aware of these potential problems
to facilitate rapid treatment of this emergency condition, potentially
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John H. Klotz is Cooperative Extension Specialist at the University of
California, Riverside. Jacob L. Pinnas and Stephen A. Klotz are Professors
of Medicine at the University of Arizona Health Sciences Center, and
Justin O. Schmidt is Director of the Southwestern Biological Institute,
American Entomologist • Volume 55, Number 3 139