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Anaphylactic Reactions

to Arthropod Bites

and Stings

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)

Phylum: Arthropoda

Class: Insecta

Order: Hymenoptera

Family: Vespidae

Genus: Vespula (ground-nesting yellow jackets)

Dolichovespula (aerial-nesting yellow jackets)

Vespa (hornets)

Polistes (paper wasps)

Family: Apidae

Genus: Apis (honey bees)

Bombus (bumble bees)

Family: Formicidae

Genus: Solenopsis (fire ants)

Pogonomyrmex (harvester ants)


Myrmecia (bulldog ants)

Pachycondyla (Asian needle and Samsum ants)

Formica (wood ants)

Order: Hemiptera

Family: Reduviidae

Genus: Triatoma (kissing bugs)

Order: Diptera

Family: Tabanidae

Genus: Chrysops (deer flies)

Tabanus (horse flies)

Family: Simuliidae (black flies)

Culicidae (mosquitoes)

Hippoboscidae (louse flies)


Genus: Glossina (tsetse flies)

Order: Lepidoptera

Family: Notodontidae

Genus: Thaumetopoea (pine processionary caterpillars)

Class: Arachnida

Order: Acari

Family: Ixodidae

Genus: Ixodes holocyclus (Australian paralysis ticks)

Ixodes pacificus (western black-legged ticks)

Ixodes ricinus


Family: Argasidae

Genus: Argas (pigeon ticks)

Order: Scorpiones

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)

Phylum: Arthropoda

Class: Insecta

Order: Hymenoptera

Family: Halictidae (sweat bees)


Genus: Pseudomyrmex (twig ants)


Rhytidoponera (green-head ants)

Order: Hemiptera

Family: Cimicidae

Genus: Cimex (bed bugs)

Order: Diptera

Family: Ceratopogonidae

Genus: Culicoides (punkies)

Family: Rhagionidae

Genus: Symphoromyia (snipe flies)

Family: Therevidae

Genus: Thereva (stiletto flies)

Order: Lepidoptera

Family: Anthelidae

Genus: Chelepteryx (white-stemmed gum moths)

Family: Megalopygidae

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

southern California

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

al. 1986).

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.

Concluding Remarks

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

saving lives. 7

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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,

Tucson, AZ.

American Entomologist • Volume 55, Number 3 139

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