Venom and the Good Life in Tarantula Hawks (Hymenoptera:

Pompilidae): How to Eat, Not be Eaten, and Live Long

Author(s): Justin O. Schmidt
Source: Journal of the Kansas Entomological Society, 77(4):402-413. 2004.
Published By: Kansas Entomological Society
DOI: http://dx.doi.org/10.2317/E-39.1
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 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
77(4), 2004, pp. 402–413

Venom and the Good Life in Tarantula Hawks (Hymenoptera:

Pompilidae): How to Eat, Not be Eaten, and Live Long
JUSTIN O. SCHMIDT
Southwestern Biological Institute, 1961 W. Brichta Dr., Tucson, Arizona 85745

ABSTRACT: Tarantula hawk wasps in the genera Pepsis and Hemipepsis are conspicuous elements
of Southwestern U. S. and the Neotropics where they often appear oblivious to potential predators
while they actively forage for nectar or search for prey. Tarantula hawks produce large quantities of
venom and their stings produce immediate, intense, excruciating short term pain in envenomed
humans. Although the instantaneous pain of a tarantula hawk sting is the greatest recorded for any
stinging insect, the venom itself lacks meaningful vertebrate toxicity. The respective lethalities of 65
and 120 mg/kg in mice for the venoms of Pepsis formosa pattoni and P. thisbe reveal that the
defensive value of stings and venom of these species is based entirely upon pain. This pain confers
near absolute protection from vertebrate predators. The pain also forms an enabling basis for the
evolution of aposematic coloration, aposematic odor, and a huge mimicry complex involving most
species of tarantula hawks and numerous flies, beetles, moths, acridid grasshoppers, and other
Hymenoptera. Tarantula hawks form mixed-species, both-sex aggregations that appear defensive in
nature and likely aid in the location of resources and mating opportunities for some species. Because
tarantula hawks have no meaningful predators, selection pressure appears to have favored long life
spans. Long-lived individuals may then function as aposematic models; thereby decreasing predatory
attacks by vertebrate predators directed toward wasp kin and future offspring. This suite of defensive
adaptations has enabled tarantula hawks to forage and behave with near impunity and to maximize
their food and reproduction while having long adult lives virtually free from predation.
KEY WORDS: Hymenoptera, Pompilidae, tarantula hawk, Pepsis, Hemipepsis, venom, mimicry

Howard Evans instilled a love of the beauty and awe of nature and science in anyone
fortunate enough to spend a few hours with him—or even in many who vicariously met him
through his books and writings. Howard loved wasps and all they could share with us, and
it showed. He profoundly affected my career and life by leading me to discover I
really wanted to be a biologist and to shed the chemistry white lab coat for an insect net,
forceps, and microscope. This all happened unbeknownst to him long before I ever met
him. His Wasp Farm (Evans, 1973) sent me hurtling down the path of investigations of
aculeate Hymenoptera and I never looked back. It is to Howard and his wasps that I dedi-
cate this paper.
Tarantula hawks (Hymenoptera: Pompilidae: Pepsinae: Pepsini: Pepsis and Hemipepsis)
are large, conspicuous, long-legged wasps whose prey are large spiders. The genera
contain several hundred mostly tropical species, with Hemipepsis distributed widely in
both Old and New World regions and Pepsis limited to the New World. In America north
of Mexico about 25 species and subspecies are known. As the common name implies,
tarantula hawks attack theraphosid spiders that are several times larger than themselves
(Petrunkevitch, 1926; Punzo, 1994b). Both male and female spiders are attacked, either in
their burrows or free ranging, and stung with rapid and complete paralysis. Paralysis is
essentially permanent, and sufficient to allow the wasp’s egg laid on the spider to hatch
and the larva to feed to completion (Petrunkevitch, 1926; Williams, 1956). Paralyzed
spiders are sometimes dragged over the surface of the ground for lengthy times and
distances as great as 100 m (Rau and Rau, 1918) before being placed in a cavity or burrow

Accepted 29 April 2004; revised 30 July 2004

Ó 2004 Kansas Entomological Society
VOLUME 77, ISSUE 4 403

and sealed. Only one spider is provisioned per larva. Female wasps sometimes dig cavities,
and sometimes use burrows of the paralyzed spider (Hurd, 1952; Williams, 1956).
With metallic black or blue-black bodies and often either bright yellow-orange or
iridescent black wings, tarantula hawks are among the most conspicuous insects on earth.
These large 3–5 cm long wasps are active during the day, and sometimes during
crepuscular times (H. E. Evans, pers. comm.), where they spend lengthy periods of time on
the ground actively searching for prey, broken by periodic slow arcing flights between
searching bouts. During the day they frequent flowers for nectar, with milkweeds
(Asclepias spp. (Asclepidaceae)) among the favorites (Hurd, 1952), and in southern
Arizona with western soapberry trees (Sapindus saponaria Linnaeus (Sapindaceae)) being
especially attractive (unpubl. data). This flashy lifestyle lends itself to risks of notice and
predation by an assortment of birds, lizards, and even mammals. Yet, very little predation
by these large vertebrate predators is noted in the literature. The answer to this apparent
paradox is the subject of this report. In keeping with the spirit I attribute to Howard Evans,
the presentation will focus on data concerning venom and other adaptive life history traits
followed by a general analysis of strategies facilitating survival and ecological success.

Materials and Methods

Wasps and Venom Collection
Wasps were collected in Southern Arizona, usually while foraging or aggregating on
nectar sources (see footnote to Table 2 for specific location information), placed in sealed
approximately 250–500 ml jars, and immediately chilled on crushed ice. Upon return to
the laboratory, the cold immobilized wasps were frozen and maintained at
248C until
venom collection. The exceptions to this procedure were wasps used for chemical analysis
of volatile odors which were dissected while cool and glands/body parts placed directly in
methylene chloride solvent (Burdick and Jackson, Muskegon, MI). Venom was expressed
from the sting tips of either cooled living, or frozen wasps as generally described in
Schmidt (1986). In brief, shortly after thawing, the entire sting apparatus was removed
from the wasp, the sting tip gently grasped about 1/4 the length from its distal tip with
Dumont No. 5 forceps, and the muscular venom reservoir gently compressed either with
blunt forceps or finger tips to express the venom through the sting shaft and by capillary
action between the tines of the forceps. Venom liquid volume was measured by collection
from the forceps into Drummond microcapillary tubes (Drummond Scientific, Broomall,
PA). The venom of some individuals, i.e., especially of P. thisbe Lucas, tended to leak
spontaneously from the sting tip upon thawing. As a result, such individuals were excluded
from the calculations for amount of venom produced per wasp. Collected venom was used
either immediately for assays, or lyophilized and stored at
248C until used.

Pain and Damage Potential of Venom

Algogenicity of stings incidentally received during field investigations was measured in
accordance with a scale of 0 to 4, with 0 representing the inability of the stinging insect to
penetrate human skin, 2 represented by the pain of a typical sting by a pollen pig (Apis
mellifera Linnaeus)(Apidae) or yellowjacket wasp (Vespula spp.)(Vespidae), and 4
representing the upper extreme as typified by stings of bullet ants (Paraponera clavata
Fabricius)(Formicidae) (Schmidt, 1990). Damage potential was measured as the lethality
of venom injected into mice as described in Schmidt (1995). Venom was used either as
fresh liquid or first lyophilized and then injected. The reason for use of liquid venom and
404 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

nearly immediate injection in some analyses was to reduce potential venom activity loss
during drying, or over time. The scarcity of available venom and the low lethality of the
venom to mice often necessitated the use of low numbers of animals for individual tests.

Life Span Investigations

Twenty two female Hemipepsis ustulata Dahlbom were collected between 13:00 and
14:30 Mountain Standard Time on the grounds of the BioSphere 2 Research Institute
located near the city of Oracle in Pinal County, Arizona on 31 December 1996. The
weather was sunny with little or no breeze and an ambient temperature of about 258C. The
wasps were foraging for honeydew produced by homopterans feeding on vigorous sprouts
from the stump of a recently cut willow (Salix sp.)(Salicaceae). The wasps were
maintained in the lab in a 36 3 50 cm (diameter 3 height) terrarium with a sandy bottom
and sticks and hiding places provided. Honey, water, and fresh powdered honeybee-
collected corbicular pollen were provided ad lib. A temperature cycle of 19/158C with
a low light level light/dark cycle of 16/8 hr was maintained. Wasps were examined daily
and those that were dead or dying removed, weighed, their venom collected, and dissected
for ovarian egg measurement.

Biological Observations
Notes from field and laboratory notebooks from 1980 to 2003 were examined for
information that might yield insights into life history strategies of Pepsis and Hemipepsis.

Results
Venom Quantities
Tarantula hawks possess long, strong, curved and sharp stingers backed with powerful
operating musculature and a prodigious amount of venom. The venom reservoir is
surrounded by thick muscle similar to that observed in vespid wasps (Schoeters et al.,
1997) which facilitates rapid venom injection into the target, be it a tarantula, a potential
predator, or an unsuspecting person. Mean expressible volumes of venom for four species
of tarantula hawks varied between 4.5 ll for Hemipepsis ustulata to 10.0 ll for Pepsis
formosa pattoni Banks (Table 1). The volume depended partially on the size of the wasp
and perhaps also on the recent venom use history for the individual. The largest amount of
venom obtained was from a large P. f. pattoni that yielded 15.0 ll. Water comprises about
67–75% of liquid venom. Average venom dry weights per wasp fell in a range of 1.35–
2.55 mg. The enormity of this quantity of venom can be appreciated when compared to the
amount produced by a single honey bee (about 150 lg), a species known to possess
relatively large quantities of toxic venom (Schmidt, 1995), or the harvester ant
Pogonomyrmex maricopa Wheeler (Formicidae) (25 lg), a large familiar stinging species
in the Southwestern U. S. (Schmidt, 1990).

Pain and Damage Potential of Venom

Few, if any, people would be stung willingly by a tarantula hawk. I know of no
examples of such bravery in the name of knowledge, for the reputation of pompilid
wasps—and tarantula hawks in specific—is well known among the biological community.
All stings experienced occurred during a collector’s enthusiasm in obtaining specimens
VOLUME 77, ISSUE 4 405

Table 1. Venom quantities in tarantula hawk wasps.

Venom volume 6 SD Venom dry Venom dry

Species (collection data)* n (range) (ll) wt. (%) wt 6 SD (mg)

Pepsis thisbe (a) 5 7.1 6 1.8 (5.0–9.1) 33.1 2.35 6 .60

P. c. chrysothemis (b) 36 5.1 6 1.4 (3.0–11.0) 27.0 1.37 6 .38
P. formosa pattoni 1982 (b) 12 10.0 6 2.6 (5.3–13.8) na 2.55 6 .67à
2000 (c) 27 10.0 6 2.0 (7.0–15.0) 25.5 2.55 6 .50
Hemipepsis ustulata (e) 6 4.5 6 1.5 (3.3–7.2) na 1.35 6 .46§

* See footnote of Table 2 for location data.

% Measured as mg dry weight of venom/ll liquid venom.
à
Using assumed venom % dry weight of 25.5.
§
Using assumed venom % dry weight of 30.0 (obtained from other samples).

and typically resulted in the stung person uttering an expletive, tossing the net into the air
and screaming—such was the immediate pain. To the author the pain was instantaneous,
electrifying, excruciating, and totally debilitating. As described by Lincecum (1867) in
reference to the effect on the prey, ‘‘the effects of the introduction of its venom is as
sudden as the snap of the electric spark’’. This description also applies to envenomed
humans. Advice I have given in speaking engagements was to ‘‘lay down and scream’’.
The reasoning being that the pain is so debilitating and excruciating that the victim is at
risk of further injury by tripping in a hole or over an object in the path and falling onto
a cactus or into a barbed wire fence. Such is the pain, that few, if any, can maintain normal
coordination or cognitive control to prevent accidental injury. Screaming is a satisfying
expression that helps reduce attention to the pain of the sting itself. Once, in his dedication
to the investigation of wasps, Howard Evans netted perhaps 10 tarantula hawk females
from a flower and enthusiastically reached into the insect net to retrieve them. Undeterred
after the first sting, he continued, receiving several more stings, until the pain was so great
he lost all of them and crawled into a ditch and just bawled his eyes out. Later, he
remarked that he made a mistake of being too greedy (A. W. Hook, pers. comm.). Not
surprisingly, the painfulness of a Pepsis sting rates 4 on the scale, the highest possible for
an insect. Among the few species that also rate a 4, a sting of a Pepsis is the most
immediately intense of all. The good news for those stung is that the pain usually decreases
to a minor level after about 3 minutes and is gone shortly thereafter, leaving the person
emotionally and physically drained. Even the immediate painfulness of the sting of the
bullet ant Paraponera clavata is less. Unfortunately, unlike the pain of a tarantula hawk
sting, the pain of a sting of that species often persists for 12 or more hours.
For mammals, tarantula hawk venom is an enormous bluff, a biological deceit of major
order. The pain speaks loudly, but no club follows. Instead of causing death or paralysis, or
leaving damage, a sting only leaves a memory in envenomed mammals. The lethality of
the venom is essentially nil. The venoms of six species of tarantula hawks were tested and
all were unremarkable (Table 2). In most cases sublethal amounts of venom caused little, if
any, effect on the envenomed animal. Even pain was not evident, except in the one
instance where an animal was accidentally injected subcutaneously instead of
intravenously with H. ustulata venom. That animal called continually for 5 min until
the venom activity ceased. Pain from tarantula hawk venom is only produced in the skin
tissues and not when introduced into the blood or the abdominal cavity. Venom challenges
as high as 128 mg/kg often were not lethal and caused minimal effect, mainly a posteriorly
splaying of the rear legs for a few minutes with the animal resuming normal appearance
406 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Table 2. Damage potential of tarantula hawk wasp venom to mice.

Venom source Venom Route LD50 (95% CI)

(collection data)* state admin n (mg/kg) Comments

Pepsis mexicana (d) liquid i.p. 1 .20 no effect

P. angustimarginata (d) liquid i.p. 1 .50 no obvious effect
P. c. chrysothemis (b) liquid i.p. 4 .61 very little effect
P. formosa pattoni #1 (b) liquid i.p. 1 ;150 died 1–4 hr; no rapid effects
#2 (f) liquid i.p. 2 3 4à 65 (29–148)
#3 (c) dried i.v. 2 .128 little effect; normal in 25 min
P. thisbe (a) dried i.v. 2 3 6à 120 (63–229)
Hemipepsis ustulata (e) dried i.v. 1 .128 legs splayed back,
normal in 5 min
dried s.c. 1 .128 constant calling for 5 min
Reference species
Apis mellifera dried i.v. 2 3 8à 2.8 (2.0–4.1) Schmidt, 1995
Vespula squamosa dried i.p. 2 3 4à 3.5 (1.8–6.1) Schmidt et al., 1980
Paraponera clavata dried i.v. 2 3 6à 1.4 (0.89–2.23) Schmidt, unpubl. data
Pogonomyrmex maricopa dried i.v. 2 3 6à .125 (0.07–0.21) Schmidt, 1990

* Location data: a ¼ AZ: Pima Co.: Tucson: SW Biol. Inst. May-June 1987; b ¼ AZ: Santa Cruz Co.: Midway
between Sonoita and Patagonia. 3–17 July 1982; c ¼ As (b). 9 July 2000; d ¼ As (b). 9–13 July 1980; e ¼ AZ:
Pinal Co.: Biosphere 2. 31 December 1996; f ¼ AZ: Cochise Co.: Portal and San Simon Valley. 25 & 29 July
1982.
i.p. ¼ intraperitoneal; i.v. ¼ intravenous; s.c. ¼ subcutaneous.
à
First number is number of challenge doses; second number is individuals per dose.

shortly thereafter. The two species for which lethalities could be determined were P. f.
pattoni at 65 mg/kg and P. thisbe at 120 mg/kg. Variability between batches of venom was
observed: for example, one batch of P. f. pattoni venom was lethal at 65 mg/kg while
another was not lethal at 120 mg/kg. Explanation for these differences is not obvious,
though in one situation fresh liquid venom was injected into the abdominal cavity and in
the other the venom was first dried and then injected into a vein. Perhaps some inactivation
of the active components occurred during the drying process, or the route of administration
is important. Future tests will be required to answer these questions. Compared to other
familiar stinging bees, wasps, and ants the venoms of Pepsis exhibit low mammalian
toxicity (Table 2). At best, tarantula hawk venom is only about 5% as lethal as that of
honey bees or the yellowjacket wasp Vespula squamosa (Drury), 2% that of a bullet ant,
and 0.2% that of a harvester ant.
Another way to measure the damage potential of an insect venom is to determine the
lethal capacity of a maximal sting: that is, the weight of a potential predator that will
receive a median lethal dose of venom if all of the venom in an individual is injected into
the animal. Lethal capacity (LC), the theoretical maximum killing power of a stinging
insect, is based on the amount of venom produced by the insect and the potency of that
venom (LC ¼ lg venom per insect/LD50). In practice the killing power of a sting would
tend to be less because not all the venom contained in the reservoir is likely to be injected
during the stinging process. The LC of tarantula hawk venom is 39 g of mouse per wasp
for P. f. pattoni but is less than 10.5 g of mouse per H. ustulata (Table 3). In contrast, the
LCs of the social bees, wasps and ants referenced in Table 3 are much greater than or equal
to that of tarantula hawks; and this is only counting one individual bee, wasp or ant. In
most defensive situations involving social insects, the potential assault against an intruder
VOLUME 77, ISSUE 4 407

Table 3. Venom lethal capacity (LC) of tarantula hawk and reference insect venoms.

Species Venom wt (mg) LD50 (mg/kg) LC (g mouse/sting)*

P. c. chrysothemis 1.37 .61 ,22

P. thisbe 2.35 120 20
P. formosa pattoni #1 2.55 65 39
#2 2.55 .128 ,20
Hemipepsis ustulata 1.35 .128 ,10.5
Reference species
Apis mellifera 0.16 2.8 57
Vespula squamosa 0.10 3.5 29
Paraponera clavata 0.30 1.4 215
Pogonomyrmex maricopa 0.025 0.125 200

* See text for further explanation.

is not simply by one individual, but by many with their additive power. In spite of their
showy appearance and enormous size, tarantula hawks cannot deliver a meaningfully
damaging sting to any but the most Lilliputian of mammals.

Life Span and Physiological Attributes of Hemipepsis ustulata

The 22 late season female Hemipepsis ustulata collected 31 December 1996 at an
elevation of approximately 1100 m appeared healthy and vigorous. They ranged in body
length from 25 to 36 mm with a mean of 31.0 6 3.2 mm. In captivity they survived on
average until 11 February 1997 (42 days after capture), with the last individual dying on
29 March. During captivity they readily consumed honey, but showed no evidence of
feeding upon the pollen that was provided. Dying wasps, when dissected contained venom
in their venom reservoirs and most contained large mature, or nearly mature, eggs. The
length of the largest egg in each wasp was 3.50 6 0.59 mm, range 2.2–4.4 mm (n ¼ 16).
Four additional individuals did not possess developed eggs. These data indicate that most
wasps were reproductively functional even during January through March.
Although not commonly seen in mid winter, female pepsine wasps are present nearly
year round in Southern Arizona. The occasional wasp is seen flying during sunny warm mid
day periods in December or January, and from captive data reported here and in Williams
(1956) H. ustulata are capable of surviving through March and are likely to be present.
Pepsis and H. ustulata appear to differ in their seasonal activity. Members of both genera
are active in spring, summer, and fall, but only Hemipepsis appears to be active during the
winter. Not known are the winter activities of Hemipepsis beyond foraging for sugar
sources. Tarantula prey are not likely to be active during this time, though we do not know
if the wasps are capable of finding tarantulas in their sealed burrows and paralyzing them.

Aggregation, Chemical Defenses, and Mimicry in Tarantula Hawks

Mixed sex and species aggregations of tarantula hawks are commonly observed. These
aggregations are sometimes loose clusters containing alert individuals with reduced
activity, and sometimes simply large active assemblages of individuals around a focal area,
often a flowering food source. Loose clusters typically occur on the shady portions of trees
during the hottest times of summer days. Active assemblages can occur throughout the day
and may be locations for foraging and mating activities. Unlike sleeping aggregations
which form during crepuscular and nocturnal times (Evans and Linsley, 1960; Linsley,
408 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

1962), tarantula hawk aggregations do not appear to be for purposes of passing the night;
rather seem designed for avoidance of poorer times during the day and potential predators.
Aggregations are sometimes encountered at night (unpubl. data; R. L. Smith, pers. comm.)
and, unlike day time aggregations, appear to be composed of mainly male individuals.
Examples of active diurnal assemblages include: 52 male and 15 female P. mexicana
(Cresson), 22 female P. chrysothemis Lucas, 15 male P. arizonica Banks, 17 male P.
cerberus Lucas, 8 male P. f. pattoni, and 5 male P. mildei Stal around an isolated blooming
Sapindus saponaria, 7 July 1980, midway between Sonoita and Patagonia, in Santa Cruz
County, Arizona; 41 male and 7 female P. f. pattoni, 13 female and 22 male P.
chrysothemis, 12 male P. mexicana, 4 male P. cerberus, 3 male P. mildei, and 2 male P.
angustimarginata Viereck, around the same tree, 10 July 1982; a mobile loose aggregation
on a mesquite tree of about 15 undetermined Pepsis that moved to stay out of the noontime
sun, 8 July 1989, Willcox, Cochise County, Arizona; and a group of 15 undetermined
Pepsis aggregating on a distal branch of a Sapindus saponaria, 11 June 1990, Tucson,
Pima County, Arizona. An important component of these aggregations appears to be
defense as indicated by the presence of a strong characteristic odor associated with the
wasps. This odor appears to the author’s nose to be the same, or very similar, for all tested
species, and both males and females produce the odor. It is released in prodigious
quantities when individuals are captured in an insect net, or when a male is held in the
hand. The source of the odor is the mandibular glands. Numerous collaborative
investigations over the last 20 years have failed to elucidate a chemical structure of the any
of the components (unpubl. data). This suggests that the active compounds are either
unstable, or very likely are compounds such as pyrazines or methyl pyrazines which can
be detected by the human nose at incredibly low levels in the air.
Tarantula hawks form a huge Müllerian and Batesian mimicry complex that includes
sexual mimicry, and mimicry by other species. Two general variations of aposematic
coloration are found in tarantula hawks: iridescent black or blue-black bodies bedecked
with bright yellow, orange, red, or brownish wings, and shiny black wings on iridescent
black or blue-black bodies. Müllerian mimicry is present among the species because
females of all species are capable of delivering formidable punishing stings to an attacker.
Males are Batesian and sexual mimics of females in both appearance and the production of
aposematic olfactory warnings. Finally, a large group of other insects mimic the behavior
and appearance of tarantula hawks (Table 4). The list in Table 4 focuses on those taxa
present in Southwestern U. S.; in the Neotropics many more taxa including arctiid moths,
tettigoniid grasshoppers, and other Hymenoptera mimic tarantula hawks (Kaye, 1913;
Evans, 1968, 1969; Evans and West-Eberhard, 1970). The mimetic relationships—
Müllerian or Batesian—of many of these species are uncertain because some including the
arctiid moths, meloid beetles, and the horse lubber T. eques (Burmeister)(Acrididae) are
chemically protected themselves.
Discussion
Tarantula hawks nearly write the manual for insect defenses. They are aposematic
brightly colored and odored, have hard, closely fitting, smooth and slippery integumentary
sclerites, live long, form defensive warning aggregations, and even have sharp and
effective defensive leg spines (J. Alcock, pers. comm.)—and they sting. These combined
properties leave little doubt that tarantula hawks are among the best defended animals on
earth. Evidence of this also comes in the form of a paucity of literature evidence for
predation on tarantula hawks; and that paucity is not for lack of interest or observation by
VOLUME 77, ISSUE 4 409

Table 4. List of some insect species in western U. S. that mimic tarantula hawk wasps.

Order: Family Species Comments: references

Excellent mimics
Diptera: Mydidae
Diptera: Asilidae
Coleoptera: Cerambycidae
Coleoptera: Meloidae
Hymenoptera: Siricidae
Hymenoptera:
Ichneumonidae
Possible mimics
Coleoptera: Meloidae
Hymenoptera: Ichneumonidae
Orthoptera: Acrididae
Mydas luteipennis (Loew)*
M. xanthopterus (Loew)*
Wyliea mydas (Brauer)
Tragidion deceptum
Hovore and Giesbert*
Lytta fulvipennis LeConte*
Urocerus californicus Norton*
Rhynchophion
flammipennis (Ashmead)
Gramptopelta sp.
Thyreodon aticolor*
flavicornis (Say)
Tegroderma aloga Skinner
Megarhyssa atrata (Fabricius)
Taeniopoda eques
Looks/behaves almost like Pepsis:
Evans and West Eberhard, 1970
Like M. luteipennis, but less bright:
Meyer et al., 1984; Nelson, 1986
Large, orange wings
Coloration/flight like Pepsis:
C. A. Olson, pers. comm.;
A. W. Hook, pers. comm.;
likely chemical defenses:
R. H. Turnbow, pers. comm.
Black with yellow/orange elytra;
movement on ground resembles Pepsis
Coloration, size, and flight
resemble Pepsis: N. M. Schiff,
pers. comm.
Black with yellow patterned wings
Black with brownish wings
Black with yellow wings
When running, raises wings exposing
red: C. A. Olson, pers. comm.
Blue-black with light yellowish wings
Males in flight resemble ‘‘pink’’ winged
Pepsis: D. W. Whitman, pers. comm.

* Additional species in the genus may also be mimics.

biologists. I am aware of no examples of amphibian, reptilian, or mammalian predation on

adult tarantula hawks. In many years of study of male and female H. ustulata, never has an
attack by a nearby bird been observed (J. Alcock, pers. comm.). Only four instances of
predation by birds have come to my attention. Wetmore (cited in Hurd, 1952) lists two
examples of kingbird predation on tarantula hawks in Puerto Rico and Punzo (1994a)
observed two instances of roadrunners attacking tarantula hawks on the ground. What is
not known is the sex of the four wasps observed being eaten, or if the two on the ground
attacked by roadrunners were healthy and vigorous at the time of attack. Roadrunner
attacks on tarantula hawks are probably uncommon at best, based on the experience of
Wade Sherbrooke, who has worked with and observed the species for years and has never
witnessed a predation (W. C. Sherbrooke, pers. comm.). What seems clear is that
predation, even on presumably harmless males, is extremely rare: such is the defensive
effectiveness of tarantula hawks.
How tarantula hawks avoid predation or injury has been the subject of many reports.
Key primary defensive properties are the venomous sting and the hard integument. In
casual conversations with biologists and the general public one is struck with the human
awe that a wasp could enter into the jaws of the largest spiders on earth and not be eaten.
Further, the impression is typically that the spider does not even try to fight back in
defense, raising the question ‘‘why not’’? As it turns out, spiders often do fight back, albeit
410 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

usually unsuccessfully. Both Petrunkevitch (1926) and Williams (1956) provide detailed
and graphic descriptions of spiders vigorously attacking and trying to crush a wasp. They
report violent encounters, often hearing loud crunching or snapping sounds as the spider
has the wasp in its jaws, and with spiders frequently losing legs during the encounters. Yet,
the wasp almost invariably escapes unharmed: no wasp deaths in 200 encounters recorded
by Petrunkevitch and only one wasp injury observed in numerous observations by
Williams. Punzo (1994a) observed that tarantulas rarely attacked their assailants, P. thisbe,
and that only one wasp death occurred in 400 encounters. These findings demonstrate
tarantula hawks do not survive simply from good fortune, but because of the amazing
defensive value of their hard, strong, slippery cuticles. Both Petrunkevitch and Williams
comment on how the crunching sounds were from the fangs of tarantulas slipping off the
smooth, hard, closely fitting integuments of the wasps.
Hard integument alone would not protect a tarantula hawk from many vertebrate
predators. The best the integument could do—and apparently does do—is to give wasps
time to deploy the sting before being damaged. The sting is their ultimate deterrence against
vertebrates. But the life history of tarantula hawks presents a conundrum: how can a wasp
have a venom that is both effective for defense against predators, yet spares the prey from
death? If a paralyzed tarantula dies before being consumed by a wasp larva, the larva dies
(Williams, 1956). The problem appears to be solved by the unusual property of tarantula
hawk venom being essentially non-toxic, but painful. Unlike ant and social bee and wasp
venoms that are usually both toxic and painful (Schmidt, 1990), tarantula hawk venom
induces pain alone in vertebrates. The non-toxic venom also has one or more unidentified
components that quickly target the neuromuscular junctions of the spider, deeply
paralyzing it. The algogenicity of the venom to vertebrates effectively repels an attack
and deters future attacks. Unlike social Hymenoptera, no queens or colonies that need
defending exist in tarantula hawks. Thus, venom lethality or damage potential is not
required to defeat the predator. All that is needed is to stop the attack and allow the wasp to
escape. Perhaps more than almost any other insect species, tarantula hawk venom has that
power: it produces the most immediate and intense pain of any hymenopteran sting. The
message perceived by humans is illustrated by the writings of Lincecum (1867) in reference
to an imagined sting to a human ‘‘I think it quite probable that the large Tarantula Killer
would produce a more serious inconvenience, and perhaps paralyze the whole system’’.
The possession of intuitively contradictory venom properties—paralysis without
lethality, and painfulness in vertebrates without either paralysis or lethality—is not
unprecedented. Different selection pressures have operated on individual scorpion venoms
to produce venom components that are selectively toxic to insects but not to crustaceans or
mammals, to crustaceans but not to mammals or insects, and to mammals but not to insects
or crustaceans (Zlotkin et al., 1972, 1975). In a similar manner, selection pressure has
resulted in individual components in black widow spider venom that are selectively toxic to
insects, mammals, and crustaceans (Fritz et al., 1980). Examples of the evolution of venom
components inducing pain without paralysis or mammalian lethality are scant and no
chemical identifications of such components in insects are known. Nevertheless, one can
envision how such a cocktail in tarantula hawk venom could evolve. Selection pressure by
vertebrate potential predators would favor individuals possessing painful venom
components, but would not favor those with irrelevant vertebrate-lethal components.
Concurrently, selection pressure against general venom lethality and subsequent loss of
prey to spoilage would be strong. These balancing forces would result in a painful, non-
toxic venom component. Prey-based selection pressure could simultaneously provide
VOLUME 77, ISSUE 4 411

strong forces for the evolution of one or more venom components that induce rapid
paralysis in the tarantula (pain should be evolutionarily irrelevant), yet are not lethal to the
spider prey. Combined competing and independent selection pressures would operate most
effectively on different venom components. Tarantula hawks evolved from an ancestor that
already possessed paralytic venoms, and from that ancestor, rapid paralysis likely evolved
first, followed by evolution of independent algogenic, non-toxic defensive components.
Complementing the primary defenses of venom and hard integument are a suite of
secondary defensive systems in tarantula hawks. The most obvious its aposematic color.
Shiny black or blue-black plus bright yellow, orange, or red are classic warning colors that
serve to remind experienced predators to avoid the bearer, or naive predators to be wary.
Enhancing the aposematic color is the aposematic odor characteristic of tarantula hawks
that, again, tips off a predator that these insects are really bad. A stung predator might well
remember the odor associated with the experience even more strongly than the color. The
combined warning system is so powerful that it serves as the basis for an extensive
mimicry complex. Not only have similar color patterns evolved in many species of
tarantula hawks, but similar colors have also evolved in a host of other organisms (Table
4). All of these organisms derive protective benefit from their convergent patterns: female
tarantula hawks benefit by having individuals of other species helping to educate potential
predators that this insect is dangerous; male tarantula hawks benefit from the protection
they gain by resembling the females; and other mimicking species gain protection from
predators via either Müllerian or Batesian mimicry of tarantula hawks. The sheer size of
the mimicry complex attests to the defensive power of the sting of the tarantula hawk.
A species that is well defended and conspicuous gains further benefit from forming
aggregations. Tarantula hawks often aggregate in localized places during times when
searches for prey are unprofitable. The presence of many individuals in an area not only
reinforces the signal conveyed by each individual, it also reduces the probability that any
given individual will be attacked, and perhaps injured, by a naive predator. In addition to
their defensive value, these aggregations appear to promote discovery of nectar feeding
locations and enhance mating opportunities. The cues used by the wasps in formation of
aggregations are not known, but the powerful residual odors produced by tarantula hawks
and their color and flight patterns are likely candidates.
When individuals are well protected, natural selection can operate in other ways to
further enhance individual reproduction and survival. One of these is increased potential
life span. The longer an individual lives, the greater its opportunities to increase its
reproduction. But also, the longer it lives, the greater its inclusive fitness. That is, by living
long and serving as a model to educate potential predators to avoid similar animals, an
individual can lower the chances that closely related individuals and future generations of
its own offspring will suffer attacks. In the case of tarantula hawks definitive information on
life spans is not known. They appear to live many months, and as shown here and by
Williams (1956), Hemipepsis ustulata can live through the winter and into spring. Although
they apparently do not live more than a year, as can occur in another painfully stinging
group, the Mutillidae (Evans and Miller, 1969; Schmidt, 1978), they live a long time by
adult insect standards. This longevity is another part of their successful defensive repertoire.
Tarantula hawks in the genera Pepsis and Hemipepsis constitute one of the most
successful groups of large insects. Their success is based largely on their defensive
adaptations which render them nearly immune to predation. The painful venomous sting is
the primary defense against large vertebrate predators and their large size plus hard
slippery integument are defenses against arthropod predators. The low risk of death from
412 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

predation, has released many constraints on their life histories and has enabled tarantula
hawks to exploit otherwise risky niches and times of day, to exhibit unimpeded and
conspicuous behaviors, to utilize dangerous prey, and to have long lives. One measure of
biological success is the ability of an organism to feed its young well, to avoid being fed
upon by other organisms, and to live a long life, usually terminated by old age. By these
standards, tarantula hawks exemplify biological success.

Acknowledgments
I thank John Alcock, Allen Hook, Carl Olson, Nathan Schiff, Wade Sherbrooke, Bob
Smith, Bob Turnbow, and Doug Whitman for providing unpublished observations and
John Alcock, Carl Olson, Nathan Schiff, and Sandy and Hayward Spangler for helpful
suggestions. Finally, I thank Mary Alice and Howard Evans who have inspired me over
the years to continue research on wasp behavior.

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