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Ann. Rev. Entomol. 1987. 32:275-95

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SCORPION BIONOMICS
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S. C. Williams
Department of Biology, San Francisco State University, San Francisco, California
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94132

INTRODUCTION

Scorpions comprise a diverse and highly successful order of Arachnids. They

have a distinctive morphology by which living and extinct members are
readily recognized. The scorpion body is conspicuously metameric with a
prosoma covered by an unsegmented carapace, a broad mesosoma consisting
of seven segments, a narrow taillike metasoma consisting of five scleroma,
and a telson modified into a stinging apparatus. The body is protected by a
sclerotized exoskeleton that efficiently retards water loss, and is set with a
variety of sensory setae and other sophisticated sensillae. Scorpions utilize an
interesting engorgement type of predatory feeding, and are adapted for surviv­
ing long periods between feedings. They use water thriftily; retention is
facilitated by excretion of guanine, uric acid, or xanthine as primary nitrogen
excretion products. Females are viviparous and, unlike most arthropods,
show maternal association with the young. Perhaps scorpions are best known
because of their venomous sting, which in some species is lethal to humans.
The antiquity of scorpions gives them a special significance. They are the
oldest known terrestrial metazoans, occurring in silurian fossils. With all the
evolutionary changes in the terrestrial environment it is remarkable that
scorpions have changed little in basic external morphology over geologic
time.
Over the past two decades the scientific community has begun to discover
the potential of scorpions as subj ects and tools of biological research, and as a
result much literature has been published. This review is limited to a summary
of recent findings in the areas of life histories, reproduction, feeding biology,
habitats, cyclic behavior, and predators.

275
0066-4170/87/0101-0275$02.00
 276 WILLIAMS

SYSTEMATICS AND DIVERSITY

Order Scorpiones is composed of eight living families: Bothriuridae, Buthi­

dae, Chactidae, Chaerilidae, Diplocentridae, Iuridae,Scorpionidae,and Vae­
jovidae. In addition, 14 extinct families are recognized. Living members of
the order show less diversity among families than insects,mites,and spiders.
Stahnke (89) estimated the numbers of described families,genera,and spe­
cies,and Williams (104) modified this estimate accounting for subsequent
taxonomic work. Francke (42) reviewed scorpion nomenclature. Today it is
estimated that there are about 1500 described living species in 112 recognized
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genera.
There are many apparently hospitable habitats in which scorpions are not
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permanently established, and also extensive habitats occupied by single spe­

cies. Single-species communities have been reported for Paruroctonus boreus
(3, 92), Vaejovis gertschi (91), and Uroctonus mordax (8). Such limited
diversity tends to occur in extreme environments,e.g. at higher latitudes and
higher elevations. In more favorable environments it is sometimes possible to
find as many as eight sympatric species. Eight species each were found in
Sonoran Desert habitats in Arizona (100) and in an intermontane basin in
Coahuila,Mexico (101). Gertsch & Allred (46) reported nine spefies from the
Nevada Test Site in southern Nevada,but this count included data from a
number of adjacent habitats, and it is unlikely that all nine species coexisted
sympatrically. Six species were reported in a California dune community
(67), five in a coastal dune habitat in Sonora,Mexico (107), and four in a
seasonally wet tropical forest in Jalisco, Mexico (105). Within fav�rable
regions three to five species are commonly encountered. Regional species
diversity of scorpions is low compared to that of spiders and mites. This low
diversity is probably due to similar habitat requirements, behavior, and
feeding biology among scorpions.

LIFE HISTORY

Life histories and population structures are hard to evaluate because of

problems in rearing litters through the life cycle and difficulties in sampling
all stadia in natural populations. The problems in determining the number of
instars in a life cycle have been analyzed (44). Life history information has
been compiled from field, laboratory,and theoretical studies. Life tables have
been constructed for Paruroctonus mesaensis (82), Urodacus abruptus (86),
and Urodacus yaschenkoi (82). All three species have high early-instar and
adult mortality,whereas intermediate instars exhibit relatively low mortality.
 SCORPION BIONOMICS 277

Life Stages
Following fertilization, young develop viviparously within the ovarian tube of
the female (41). Young are born in an early stage of development called the
larva (first instar of life cycle). The larval stage differs from the subsequent
stages in that larvae lack ungues, a fully formed telson, and a fully sclerotized
exoskeleton; they also differ in their response to the environment, in part
because of their smaller surface-to-volume ratio. They do not feed, are not
capable of significant independent locomotion, and cannot sting. They nor­
mally assume a position on the mother's dorsum and molt to the second instar
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stage (first nymphal instar) in about 1-2 wk. Scorpions of most species
disperse from the mother as second instars; in some species this stage may be
the primary dispersal stage. Nymphs molt several times before reaching
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sexual maturity. Scorpions of the few taxa studied have 5-7 stadia. The
number of instars in the life cycle (including larva and imago) has been
estimated at seven in Uroctonus mordax (37), Paruroctonus mesaensis (70),
and Centruroides gracilis (43) and six in Buthotus minax (90), Urodacus
abruptus (86), and Urodacus yaschenkoi (83). Some species may mature in
more than one stadium. Some males of Tityus trivittatusJasciolatus mature in
the fifth instar, while others mature in the sixth (57, cited in 44). Females,
however, appeared to all mature as sixth instars. A similar pattern of males
maturing in more than one instar has been reported for several species (44). It
is believed that scorpions do not molt after reaching sexual maturity, but they
may live for several years in the adult stadium.

Growth Rate

Many species reach maturity during the first year of life. This has been
determined for C. gracilis (41). However some, such as P. mesaensis, have
been estimated to mature in 19-24 mo (72), and U. abruptus requires 3 yr
(86). In a tabulation of the time required for 10 species to reach sexual
maturity (43), Buthotus minax was the fastest, reaching sexual maturity in
164-307 days, while Tityus trivittatus was the slowest, requiring 426-810
days.
Most scorpions experience the first molt within 2 wk, and the following
five or six molts occur at various intervals during the next 9-12 mo. Growth,
as a result of molting, is in linear dimensions only, because mass decreases
slightly as a result of ecdysis. Comparison of postmolt to premolt lengths has
Uroctonus mordax, 20-45% (37); Isometrus
revealed the following increases:
maculatus, 30% (74); Buthus occitanus, 26% (7); and Tityus bahiensis, 22%
(62, cited in 44).
 278 WILLIAMS

Longevity
Scorpions vary considerably in longevity. In a number of taxa females tend to
live longer than males; mature males may have a higher mortality rate because
of increased exposure to predation and suboptimum habitat conditions during
the mate-seeking season. Estimates of maximum longevity for some species
are as follows: Paruroctonus mesaensis, 5 yr (68); Urodacus abruptus, 10 yr
(86); Centruroides gracilis, 4 yr (43); Centruroides sculpturatus, longer than
5 yr (88); u. mordax, longer than 579 days (37); Tityus bahiensis, 1417 days
(62, cited in 44); and Tityus serrulatus, 1365-1565 days (64).
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REPRODUCTIVE BIOLOGY
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Courtship and Mating

Mating is preceded by an elaborate courtship, referred to as the "promenade a
deux," which has similar behavioral components throughout the order. Court­
ship and insemination normally occur during the night on exposed habitat
surfaces. Fertilization is internal. Males, however, lack a penis and depend on
a spermatophore for sperm transfer. The courtship is a curious behavioral
sequence during which mates locate and orient to each other. They grip each
other by the pedipalp chelae and sometimes also by the chelicerae, and walk
forward and backward in coordinated movements that resemble a dance until
a suitable substrate for spermatophore deposition is found. The male emits an
elongated spermatophore, which is usually glued to a firm surface; he then
pulls the female over the spermatophore, and insemination takes place.
Courtship has been described numerous times in a variety of taxa (9, 10,
84, 87, 88, 102). The use of spermatophores was not detected until 1955 (1,
2, 4-6). It now appears that male scorpions universally inseminate females
indirectly in this way. The uptake of sperm by courted females may not
always be successful (2).
Polis & Farley (71) described an episode of courtship and insemination in
Paruroctonus mesaensis. The initial contact phase of courtship lasted about
22 min and involved physical contact, movement, and a distinct vibration of
the body called "juddering." Following initial contact the pair went into the
courtship dance, which lasted 9.3 min and ended in spermatophore deposition
and insemination. During the promenade the pair traveled approximately 8 m.
Insemination took 3-5 sec.
Parthenogenesis is known in scorpions but is not common. It has been
reported in Tityus serrulatus (61, 63, 64, 79) and in Liocheles australasiae
(59, 60). In L. australasiae thelytokous parthenogenesis was observed in the
laboratory, and field observations suggest that some, if not most, populations
of this species reproduce only in this fashion.
 SCORPION BIONOMICS 279

Gestation
Many temperate taxa, such as members of Vaejovis, Uroetonus, Hadrurus,
Serradigitus, and Paruroetonus, appear to normally have one litter per year,
with births more or less synchronized within a given population. Some more
tropical taxa, such as members of Centruroides, may have one to several
litters per year, and data suggest that gestation intervals are 2-4 mo. The
gestation period for a number of species has been estimated as follows:
Centruroides insulanus, 4.5-7.5 mo (10); Centruroides vittatus, 8 mo (10);
Urodaeus abruptus, 16 mo (86); and Paruroetonus mesaensis, 1 yr (72).
Probable causes for variation in gestation among species include size, number
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of young born, and general climatic conditions experienced by the population

(102).
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Scorpions are categorized as apoikogenic and katoikogenic based on the

nature of their internal development (41). In apoikogenic forms the embryos
are each covered by a birth membrane formed by the fusion of the embryonic
serosa and amnion. In contrast, katoikogenic embryos are not covered by
extraembryonic membranes; they develop in large diverticulae opening into
the reproductive tube. Katoikogenic forms are consistently born tail first,
while apoikogenic forms are born either head or tail first.

Parturition

The birth process has been described a number of times and for a variety of
taxa (40, 41,52,85, 102). The mother stilts high above the substrate on her
hind two pairs of walking legs; her genital opercula open and the young
emerge, one by one, through the genital aperture. In most families, each
young is covered at birth by a birth membrane. In contrast, members of the
Diplocentridae and Scorpionidae are not covered by such membranes (41). As
the young emerge, the mother usually catches them in a "birth basket" formed
by the first two pairs of walking legs. In the birth basket, young that are
enclosed in birth membranes shed them. Then they climb the mother's
anterior walking legs to her dorsum. As larvae ascend to the mother's dorsum
they may assume either an ordered orientation (Vaejovis) or, more commonly,
a random orientation (Centruroides, Uroetonus, Syntropis, and Megacormus)
(37, 38, 51, 52, 102). The significance of the highly ordered orientation in
Vaejovis is still not known.
The length of parturition varies among individuals of the same species and
among members of different species. In general, the time spent in parturition
depends on the number of young delivered, the size of young, and the
occurrence of complications. One Hadrurus arizonensis female delivered 10
young in 1 hr, while one Vaejovis spinigerus delivered 69 young in 7.5 hr
(102). Apoikogenic forms tend to have more rapid parturition, ranging from
 280 WILLIAMS

55 min to 37 hr, while parturition for katoikogenic forms ranged from 36 to 74

hr (41).
Parturition is a critical point in the life cycle; if the young are unable to shed
their birth membranes or ascend to the mother's dorsum their probability of
survival is low. In U. abruptus the mortality during birth is about 27% (86).

Maternal Care
Once on the mother's dorsum larvae assume a characteristic posture and
remain there throughout the ftrst stadium. Within 2 wk the larvae syn­
chronously molt to the second instar stage (the ftrst nymphal instar) and
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remain on the mother's dorsum for 1-2 wk, at which time they begin
dispersing. In some genera, such as Vaejovis, Uroctonus, Paruroctonus, and
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Hadrurus, mothers remain inactive within their protective shelters while with
young. In others, such as Centruroides, the females may be seen during
nocturnal hours in exposed areas with young on their dorsum.
During the ftrst stadium there is no obvious nutrient exchange between
mother and offspring. Second instars have been observed feeding on remains
of their mother's prey, but they generally begin dispersal in this stadium, soon
after their cuticle hardens. Thus, maternal feeding of young is probably
insigniftcant. In a contrasting observation, however, a Palamnaeus longima­
nus mother provided food for her litter,which was reported to disperse in the
third stadium (81). Vannini and coworkers (95) recently investigated the
question of nutrient exchange between mother and larvae. In experiments
with Euscorpiusjlavicaudis, mothers given tritiated water transferred tritium
to larvae on their dorsum. Tritium may have been incorporated in epicuticular
lipids which were then transferred to the larvae by contact; or it may have
been absorbed by the larvae in the form of liquid water or vapor from the
mother. This study suggests that a reexamination of the adaptive scope of
mother-larval association is in order. The mother-larval association was
recently investigated in three species of Euscorpius (96-98). Larvae were
observed to mount substitute mothers, and in the absence of a mother larvae
mounted artificial models and dead scorpions, but they did not settle on these
permanently. Larvae settled permanently on the mother's dorsum owing to
substratum recognition. Chemical recognition was not species-specific,nor
did it carry information on sex or reproductive phase. Three adaptive benefits
of mother-larval association have been hypothesized: antipredator defense;
continuous maternal selection of the optimal microclimate for larvae; and
trophic exchange between mother and larvae (97).

Fecundity
The number of young in a litter varies. Scorpions have been reported to have
6-105 young per litter. Litter sizes have been reported as follows: Centrur-
 SCORPION BIONOMICS 281

oides insulanus, 20-105 (10); Centruroides sculpturatus, 12-36 (88); Cen­

truroides vittatus, 20-47 (10); Diplocentrus bigbendensis, 37 (40); Dip­
locentrus spitzeri, 9 (39); Hadrurus arizonensis, 10 (102); Megacormus
gertschi, 19-75 (38); Paruroctonus boreus, average of 34 (92); Paruroctonus
mesaensis, 18-21 (68); Uroctonus mordax, 28-34 (37); Urodacus abruptus,
9-17 (86); and Vaejovis spinigerus, 66-69 (88, 102). Litter size and number
of litters produced per female are perhaps related to species group, population
density, and availability of prey; however, the influence of these factors has
not been established.
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FEEDING BIOLOGY
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Scorpions are essentially obligate predators. They seldom ingest dead prey.
However, adults occasionally accept freshly killed prey in the laboratory.

Prey Encounter
The several strategies that scorpions have evolved to encounter prey can be
put in three primary classes: waiting on exposed substrates, active stalking,
and waiting in their burrows. The most common strategy is to leave the
diurnal shelter in early evening, settle in an exposed location on the ground
surface or other exposed substrate, and wait for prey. Prey is captured only if
it encounters the scorpion or comes close enough to stimulate the sensory
receptors of the quiescent scorpion. Most species of Vaejovis,Paruroctonus,
Vejovoidus, Uroctonus, Serradigitus, and Hadrurus use this strategy. Mem­
bers of Centruroides, in contrast, sometimes actively traverse exposed sub­
strates, apparently searching for prey. Members of Anuroctonus phaiodacty­
Ius use their burrows to trap prey. The scorpion remains quiescent just inside
the entrance to its burrow and captures insects that enter. Examination of the
remains of prey in the burrow tumuli has suggested that these scorpions may
be diurnal predators. Superstitionia donensis has been observed living in a
shallow cell under cattle dung, where it preys on dung-feeding insects.
Studies have revealed that Paruroctonus mesaensis has a sophisticated ability
to locate and orient toward prey using prey-generated low-frequency com­
pressional vibrations in sand (12-15). It uses basitarsal slit sensillae to detect
these vibrations, and can effectively locate moving prey at a distance of up to
50 cm. In P. mesaensis this appears to be the primary means of locating prey.
Even prey burrowing under the sand surface are detected and caught by a
thrust with the pedipalps. A similar pattern of orientation has been observed in
Hadrurus arizonensis, but the range and accuracy of this scorpion's response
to prey was less than that observed for P. mesaensis (15). Scorpions have
other sense organs (mechanoreceptive setae, chemoreceptive setae, and che­
moreceptive tarsal organs) that may aid in predation (36).
 282 WILLIAMS

Prey Capture
Scorpions firmly grasp encountered prey with the chelae of the pedipalps.
Some scorpions, such as Centruroides exilicauda, then sting the prey before
ingestion. Others, such as Hadrurus arizonensis and Anuroctonus phaiodac­
tylus, often do not attempt to sting the prey although they are capable of doing
so. When the sting is not used or is not lethal, the scorpion firmly immobilizes
its prey with the pedipalp chelae and ingests it alive. Some species have
venoms that show no effect on prey. Scorpions probably kill their prey more
often by eating them alive than by injecting venom.
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Scorpions engorge when prey are available, and may remain for a long time
in a quiescent state between feedings. Feeding scorpions ingest a high propor­
tion of their prey's biomass, may increase their own biomass substantially
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through engorgement, and utilize a preoral exudate that appears to initiate

digestion in the carcass of the prey before ingestion. Vaejovis gertschi in­
creased body weight by 9.6-16.5% during a single feeding (91). Weight
increases of 9-33% in A. phaiodactylus and 10-36% in U. mordax have been
measured (35).

Ingestion
Following immobilization of prey, the scorpion uses its chelicerae to tear it
open, and pumps a preoral exudate from the scorpion's buccal region into the
carcass. A rhythmic flow of this fluid from the scorpion to the carcass and
back is established shortly and lasts throughout ingestion. AnuroctoTluS
phaiodactylus fed on crickets ingested 8-97% of the prey biomass, and
UroctoTlUS mordax showed similar ingestion efficiency of 5-87% (35). The
efficiency of ingestion is striking, and is probably due to digestive enzymes in
the preoral exudate, a setal mechanism in the cheliceral region that minimizes
fluid loss, and an efficient gnathobase press that separates fluids from particu­
late matter. During the evening, under natural conditions, scorpions ingest
prey on their habitat surface or a! burrow entrances. Most prey of observed
Paruroctonus mesaensis were eaten at the capture site, but about one quarter
were carried to nearby vegetation for ingestion (67).

Predation Frequency
Observations suggest that scorpions normally capture and ingest prey in­
frequently, and that they are well adapted for intermittent feeding. Vaejovis
gertschi feeding on termites lost weight in a linear fashion after engorgement
(91). On the average these scorpions returned to their prefeeding weights in
13 days; this suggests that individuals need to feed about once every two
weeks to maintain their biomass (at 22°e and near 100% RH) (91). In a
similar study, Anuroctonus phaiodactylus individuals feeding on crickets
returned to their prefeeding weights in 10-23 days, whileUroctonus mordax
individuals returned to their prefeeding weights in 5-68 days (35); individuals
 SCORPION BIONOMICS 283

of these species apparently need to feed once every one to ten weeks to
maintain nonnal biomass. Under natural environmental conditions individual
scorpions need to feed only periodically; the periodicity is probably de­
tennined by size of prey, temperature, relative humidity, and physical activity
of the scorpion.

Prey
Prey selectivity by scorpions has been reviewed(8-10, 35, 65, 67, 87, 88,
91, 92). Scorpions will capture almost any prey encountered that they can
physically immobilize and ingest. They feed primarily on insects, arachnids,
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and other arthropods. Predation on mollusks has also been reported (55) but is
not common. Some larger species also prey on small vertebrates. In one study
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Vaejovis gertschi accepted geometrid caterpillars, ichneumonids, silverfish,

machilids, tipulids, termites, and ground-dwelling spiders as prey (91), but
rejected crickets, small roaches, mirids, carabids, tenebrionids, and mil­
lipedes. Vaejovis confusus. Vaejovis spinigerus. and Hadrurus arizonensis
prey on a wide variety of arthropods (100). Hadrurus arizonensis is dis­
tinctive in that it readily eats lizards and mice. Paruroctonus mesaensis also
preys on a variety of arthropods, and moreover shows a substantial tendency
toward cannibalism (50, 67). In a California dune community 9% of the diet
of P. mesaensis was estimated to be supplied by cannibalism (70). This
impressive level of cannibalism was not detected in a different study on this
species in another area(50). Polis (67) recorded 95 prey species accepted by
P. mesaensis 81 of these comprised only 1 % of the
in a sand dune habitat;
scorpion's diet. He reported that 10% of the prey were fossorial, 80% were
cursorial, and 10% were aerial. The major prey composition was 42% tenebri­
onids, 17% orthopterans, 16% other scorpions, and 12% hymenopterans.
Wonn snakes (Leptotyphlops) were also eaten. Urodacus abruptus has been
reported to accept a variety of prey, mainly insects and spiders; they exhibited
no sign of cannibalism or aggressive intraspecific behavior when they were
well fed (87).
Scorpion predation on vertebrates has been summarized (65). Vertebrate
prey has included 2 species of mammals, 11 species of lizards, and 2 species
of snakes. Ten species of scorpions have been reported to prey on vertebrates :
Pandinus pallidus. Opisthophthalmus carinatus. Hadogenes sp., Parabuthus
villosus, Vaejovis sp., Paruroctonus mesaensis, Hadrurus arizonensis, Had­
rurus spadix, and Hadrurus hirsutus.

HABITATS

Scorpions occupy a greater range of habitats than is generally recognized.

They are most abundant and diverse in arid environments of lower temperate
latitudes. The most diverse assemblage of scorpions known is in Baja Califor-
 284 WILLIAMS

nia, Mexico, where 61 species, 11 genera, and 5 families have been reported
in the narrow peninsula (104). In this region scorpions are found in stabilized
and unstabilized sand dunes, littoral wrack and cobblestones, basaltic cliffs,
rock slides, burrows in soil, rock crevices, mud cracks, vegetation, and
skeletons of dead cacti; and under stones, bark of trees, surface debris, and
cattle dung. Some species, particularly members of the genus Centruroides,
are noted for adapting to human habitation, where they may be numerous in
roof thatching, between bricks and stones in walls, in firewood, and in yard
debris. In moist tropical forests of Mexico scorpions are found in the forest
canopy, in bromeliads, and on tree trunks. Some of these scorpions, such as
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Centruroides thorelli, complete their life cycle in the forest canopy and
perhaps never descend to the ground. Some scorpions are also adapted for
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cave existence (58, 66).

Scorpions are generally thought of as inhabitants of xeric environments;
indeed some fonns, such as Serradigitus deserticola, are found in habitats
that may only receive rainfall every few years. At the other extreme, Urocto­
nus mordax may be found under damp mosses in moist habitats of northern
California.
Scorpions have generally been considered dwellers of low elevations, but
populations have recently been detected at elevations of 2133 m in the Sierra
Nevada of California (106). The range of scorpions extends down to sea level,
where species like Vaejovis littoralis live in the littoral wrack zone along the
shore; Centruroides exilicauda and Vaejovis janssi are active predators along
beaches (104). Some scorpions have established populations that extend to
high latitudes. In North America, Paruroctonus boreus is found to about 5r
N latitude in British Columbia, Alberta, and Saskatchewan (16, 47).
Even though some species are widely distributed, there is evidence of
habitat preference and a habitat-related limitation of distribution. Allred (3)
reported habitat preferences among 283 P. boreus individuals collected in 8 of
12 different plant communities in Idaho. The ideal habitats were those
vegetated with 85% ground cover composed of at least 60% broadleaf shrubs
and 5% grass. In a similar study in southern Nevada, P. boreus was most
abundant in pinyon pine and juniper communities (46). This species and most
individuals of eight others were broadly distributed among several habitat
types, but also showed preferences. Vaejovis gertschi has shown preference
for three natural habitats in central California: open rock surface, rock crev­
ice, and vegetation-covered soil (91). The habitat preference changed sea­
sonally. The main population migrated to elevated rock outcrops during the
winter (rainy season). In spring they migrated back to the adjacent grassland,
where they used fissures in the drying soils as diurnal shelters. Vaejovis
confusus and Vaejovis spinigerus both occurred in the same region of the
Sonoran desert, and although their distributions overlapped, their different
 SCORPION BIONOMICS 285

microhabitat preferences allowed them to avoid competitive interactions

(103). Habitat preferences have also been reported in the Australian species
Urodacus abruptus (86).

Burrowing Biology
Burrowing is a major adaptation of many scorpion species for survival in
extreme environments (48, 99). Burrows may be as simple as a cell excavated
under rocks, logs, or other surface cover, or they may be elaborate and deep.
Paruroctonus mesaensis and Vejovoidus longiunguis dig relatively shallow,
temporary burrows, while other scorpions such as Anuroctonus phaiodactylus
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construct permanent burrows, sometimes to depths of up to 42 cm (99).

All 19 species of the African genus Opisthophthalmus burrow in the ground
for shelter (56). Lamoral (56) found that habitat selection, distribution, and
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range of each of these species were correlated with particular soil types. The
most important soil factor was hardness and texture. Differential interspecific
soil-hardness requirements among burrowing species of Opisthophthalmus
insured that different species in the same area were ecologically allopatric,
and thus prevented competition for burrowing sites.
The structure of burrows has been related to their functions. Koch (54)
found significant structural differences in burrows of Urodacus hoplurus and
Urodacus yaschenkoi. Both of these Australian species resemble the North
American Anuroctonus phaiodactylus in that they construct burrows with a
distinct entrance followed by a tunnel (tortuous and spiral in Urodacus,
usually simpler in Anuroctonus) and a terminal chamber. Urodacus yaschenk­
oi excavated normal burrows within 8-10 hr in the field (83). The burrows
were maintained throughout the year; the most obvious maintenance was
clearing of debris from the entrance. Shorthouse & Marples (83) concluded
that this burrow structure is adaptive because vertically spiralling tunnels are
less likely to intersect adjacent burrows than are oblique straight tunnels; the
entrance chamber facilitates predation; and the deep terminal chamber may
facilitate avoidance of environmental extremes. Burrowing behavior results in
avoidance of high daytime temperatures and low humidities, minimization of
water loss, and probably lowering of metabolic needs (82).
The subterranean environment of burrowing scorpions differs from the
surface environment. Crawford & Riddle (24) recognized that the environ­
ment of the burrowing Diplocentrus spitzeri is essentially mesic, even though
the ground surface is xeric. This species frequently constructs its burrow
entrance under a surface rock, which gives additional protection to the
burrow. These authors suggested that D. spitzeri could survive in the absence
of free water if sufficient prey were available; thus drought should not
normally cause significant mortality for individuals in established burrows.
They also concluded that high temperature does not usually cause significant
 286 WILLIAMS

mortality because of the tempering effect of the burrow, but that exposure to
extreme winter cold could occasionally result in mortality.

Home Range
A few species, such as Anuroctonus phaiodactylus and Didymocentrus com­
ondae, appear to be restricted to their burrows throughout their lives. These
species have home ranges limited to the burrow and a modest surface area
surrounding the burrow entrance. Paruroctonus mesaensis usually remains
within 0. 5 m of its burrow entrance at night (50). Tourtlotte (92), in a
mark-recapture study of Paruroctonus boreus, noted individual recaptures in
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approximately the same locations in two successive years.

Dispersal
by University of Edinburgh on 11/02/10. For personal use only.

Scorpions probably disperse less than is generally believed. The suggestion

that scorpions are transient by nature has come from frequent observations of
scorpions entering human habitation. Most of those observed have been
mature males that have become more ambulatory during courtship season.
Williams (100) found that most juvenile stadia of Vaejovis spinigerus, Vae­
jovis confusus, and Hadrurus arizonensis were not commonly encountered on
the habitat surface in the Sonoran desert. Dispersal was primarily accom­
plished by nymphs in the first postiarval instar and adults, especially males.
Some island scorpions disperse by rafting on logs and other floating debris. In
the Gulf of California, rafting species such as Centruroides exilicauda are
widely distributed among the numerous islands, while nonrafting species,
such as members of Vaejovis, have more restricted distributions and more
extensive allopatric speciation (104).
Some scorpions are dispersed by humans. The large tropical American
species Centruroides gracilis is commonly found alive in imported produce
and forest products at ports of entry throughout the world. Over the past
several decades campers from the Colorado River region have repeatedly
introduced Centruroides exilicauda into southwestern California, and there is
now evidence that it has become established there (45, 78).

CYCLES

Scorpions exhibit conspicuous activity cycles. In captivity, and probably

under natural conditions as well, they spend most of their time in an immobile
state. During these periods of suppressed activity their sensory awareness is
also depressed. Periodically, however, scorpions become physicaIIy active
and engage in predation, burrowing, and courtship. In the inactive state,
scorpions are usually located in their protective shelters. During the active
phase of their cycle they may come to the surface. Perhaps their extensive
 SCORPION BIONOMICS 287

inactivity is related to their dependence on relatively inefficient booklungs for

their oxygen supply. The oxygen consumption of scorpions suggests that their
respiratory rates are among the lowest known for terrestrial animals (24-27,
49, 76-77, 93). Their minimal physical activity, coupled with low energy
requirements, is an adaptive strategy for survival in unfavorable or highly
cyclic environments. In the discussion that follows, the term "surface activ­
ity" is used to indicate movement from the shelter to the exposed surface of
the environment, even if this does not entail physical activity on the surface.
On any given night only a small subset of a scorpion population surfaces,
and the proportion of the population that surfaces is influenced by environ­
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mental parameters such as temperature, rainfall, relative humidity, moon­

light, and season. In New Mexico, about 5% of a population of Paruroctonus
utahensis came to the surface on any night during a favorable season (11).
by University of Edinburgh on 11/02/10. For personal use only.

About 15% of a population of Uroctonus mordax studied in central California

between May and September was active on the surface at any time (including
those in burrow entrances) (8). About half of a P. mesaensis population was
on the surface of a southern California sand dune during a given evening (69).

Seasonal Cycles
Activity cycles of scorpions are correlated with seasonal changes. In Idaho,
Paruroctonus boreus was observed on the ground surface only from March
through October, with highest surface densities in July and August (92).
Similar seasonal patterns in surface activity have been reported in An­
uroctonus phaiodactylus (46, 99), Paruroctonus becki (46), Paruroctonus
boreus (3, 46), Paruroctonus mesaensis (69), Paruroctonus utahensis (11,
77), Uroctonus mordax (8), Vaejovis confusus (46, 100), Vaejovis gertschi
(91), Vaejovis hirsuticauda (46), Vaejovis spinigerus (100), Vaejovis wupat­
kiensis (46), and Hadrurus arizonensis (46). In the western United States
scorpions can generally be observed on the surface from March through
October, with highest surface densities from May through September.
Seasonal surface-activity patterns may be influenced by temperature, rain­
fall, courtship season, and moonlight. Positive correlations between tempera­
ture and surface activity have been reported in Paruroctonus boreus (92),
Uroctonus mordax (8), and Vaejovis gertschi (91). Critical temperatures
below which scorpions do not ordinarily surface have been identified for a
few species: P. boreus, IDoC (92); P. mesaensis, adults 20°C, juveniles 9°C
(69); U. mordax, 4°C (8); and V. gertschi, 4°C (91).
Precipitation also generally affects surfacing. During and immediately after
precipitation, surface densities of P. mesaensis were reduced (69). P. boreus
exhibited no surface activity during precipitation, and the surface densities
remained reduced for 3-5 days following substantial precipitation (92). The
surfacing of V. gertschi was negatively correlated with relative humidity (91).
 288 WILLIAMS

Illumination influences the surfacing of some species. The surface densities

of P. mesaensis (50), U. mordax (8), V. confusus (50, 100), V. gertschi (91),
and V. spinigerus (100) appear to be negatively correlated with intensity of
moonlight. By contrast, surfacing of C. sculpturatus (50) and P. mesaensis
(69) was reported to be uncorrelated with moonlight. P. mesaensis was
inhibited by moonlight in a coastal dune community but not in an inland dune
community; this suggests a difference in adaptation to predation pressures.
Ninety-five percent of observed matings by P. mesaensis occurred on moon­
less nights (69). During the full moon only 1.6% of surfaced individuals of
this species were feeding, in contrast to 3.8% during the new moon.
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Surface activity usually increases significantly during courtship season,

particularly among mature males. Moreover, males appear to abandon their
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normal burrows or other shelters and become more nomadic, and thus show
increased surface locomotion. This results in observed sex ratios that strongly
favor males. The influence of courtship on surface activities is evident for
Anuroctonus phaiodactylus (99), Uroctonus mordax (8), Paruroctonus
boreus (3, 46, 92), P. mesaensis (69), Vaejovis confusus (100), V. gertschi
(91), and V. spinigerus (l00).
Scorpions generally remain relatively inactive following a feeding to satia­
tion. Bradley (11) found that when Paruroctonus utahensis had fed, they did
not return to the ground surface for an average of 16-20 days. He concluded
that they remained in their protective burrows to minimize exposure to
predation.

Circadian Cycles
Some of the most obvious activity cycles exhibited by scorpions are circadian.
Most species exhibit such cycles. Most are considered nocturnal, in that they
reside in their shelters during the day and come to the surface after sunset. In
contrast, a few species display primarily diurnal activity. Vaejovis littoralis, a
coastal littoral dweller, has activity cycles correlated with tidal movement,
and is active during the incoming tide, even during midday (104). Circadian
surface activity has been reported in the forest-dwelling Pandinus imperator
(diurnal) and in Buthus hottentotta (nocturnal) (94). Centruroides sculptura­
tus and Diplocentrus spitzeri both have nocturnal activity under a normal
photoperiod, but when kept in constant darkness C. sculpturatus displayed an
endogenous circadian rhythm, while D. spitzeri did not (23). The contrasting
circadian rhythms of these two species may create a temporal separation that
ultimately facilitates sympatric coexistence (23). Endogenous circadian
rhythms in physical activity have also been reported in a number of other
species (17-22).
Nocturnal species do not come to the surface every night. A periodic
pattern of surfacing has been reported in several species. Those species
 SCORPION BIONOMICS 289

studied characteristically surfaced shortly after sunset and remained on the

surface for varying periods. Surface density is usually highest during the first
few hours following sunset. Surfaced scorpions gradually return to their
shelters throughout the evening, and by midnight a large proportion have done
so. By sunrise few if any normally remain on the surface. This pattern has
been observed in a number of species (23, 50, 70, 82, 91, 94).
Enzymatic and neurophysiological functions also have circadian rhythms
and are correlated with the activity cycle of scorpions (48). Muscle de­
hydrogenase activity in Heterometrus julvipes is highest at times of highest
locomotion and oxygen consumption (75). Circadian rhythms are found in the
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phosphorylase activity and glycogen content of the heart muscle of H. fulvipes

(53). The neurophysiological basis of circadian rhythms has been studied in
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the North African species Androctonus australis and four other species (29-
31, 33, 34). These studies reveal that light sensitivity of the ocelli is con­
trolled by a circadian clock located in the central nervous system. The median
eyes were found to be 1000 times more sensitive during the subjective night
than during the subjective day. Efferent neurosecretory fibers mediate the
circadian signal from the central nervous system. The sensitivity cycle of
median and lateral ocelli is synchronized, but the median ocelli are far more
responsive to the sensitivity cycle. This circadian rhythm is synchronized to
varying environmental conditions by an external zeitgeber (28, 31, 33). Both
median and lateral ocelli are capable of mediating zeitgeber stimuli, but
lateral ocelli are more sensitive in this function (31, 32). The lateral ocelli
sacrifice image information in favor of increased sensitivity, and show small
fluctuations in sensitivity (80). The combination of both types of ocelli is
believed to enable the central nervous system to calculate the time of dusk and
dawn more precisely than it could with input from either type alone (28, 29).
Electroretinograms from five species of Buthidae and Scorpionidae suggest
that use of ocelli and zeitgeber stimuli may be a general mechanism for
regulating and synchronizing circadian rhythms in scorpions. Extraocular
light reception through the exoskeleton of the metasoma has been observed in
a species of Urodacus (108); these scorpions were able to orient to dark and
light habitats exclusively by this means (l09).

PREDATORS OF SCORPIONS

Scorpions are potentially attractive prey. Factors contributing to their de­

sirability as prey include large body size,rich nutrient content, abundant
populations, wide distribution,relative lack of defense,and predictable sur­
face behavior. At least 124 vertebrates and 26 invertebrates prey on scorpions
(73). These data suggest that vertebrates are more important than in­
vertebrates as scorpion predators.
 290 WILLIAMS

North American vertebrates that may utilize scorpions as a major part of

their diet include elf owls (Micranthene), burrowing owls (Speotyto), barn
owls (Tyto), grasshopper mice (Onychomys), desert shrews (Notiosorex), and
a variety of lizards. Some of these protect themselves from scorpion sting by
removing the telson at the time of capture. This is a curious behavior, since
the venom of most scorpions appears to have little if any effect on vertebrate
predators.
The importance of invertebrates as scorpion predators has been un­
derestimated because (a) they have been recorded as predators only when
actually observed preying on scorpions; (b) invertebrate stomach contents
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have not been evaluated; (c) invertebrate predators do not usually leave
conspicuous remains similar to owl pellets, scats, or unconsumed remains;
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and (d) after scorpions have fed to satiation and have returned to a catatonic
state in their shelter, they may fall prey to small insects that otherwise might
not prey on them. In the laboratory, inactive scorpions are commonly killed
by meal worms, crickets, and other small insects supplied as prey. Black
widow spiders (Latrodectus), Jerusalem crickets (Stenopelmatus), darkling
beetles (Eleodes), centipedes (Scolopendra), and solpugids are known to feed
on scorpions.
In a California dune community, 11 species of scorpion predators were
identified (67). These included loggerhead shrikes, great homed owls, kit
foxes, grasshopper mice, harvester ants, black widow spiders, and five
species of scorpion (70). Extensive predation on scorpions by other scorpions
was observed.

CONCLUSIONS

There has been much activity in scorpion research, yet studies have been
based on relatively few species. Of the estimated 1500 species, only three to
four dozen have been studied extensively. The results of recently introduced
ultraviolet sampling suggest that our knowledge of the composition and
distribution of the scorpion fauna is far from complete. Recent field studies
indicate that many new taxa remain to be discovered. It is hoped that
discovery of new taxa will stimulate a much-needed reinvestigation of the
systematic and biogeographic relationships among scorpions. Scorpions
should continue to be an important tool in the study of circadian rhythms and
their neurological bases. The importance of scorpions as predators and
stabilizers within terrestrial COfnmunities is still not fully appreciated. The
impressive adaptations of scorpions to rigorous environments continue to
attract the attention of physiologists, behaviorists, and ecologists, and our
understanding of these adaptations is certain to be significantly enhanced.
Scorpions will most likely become even more popular in research as their
taxonomy becomes better known.
 SCORPION BIONOMICS 291

ACKNOWLEDGMENTS
Much appreciation is due my colleagues Paul H. Arnaud, Jr., Jack T.
Tomlinson, and John E. Hafernik for kindly reading and criticizing this
manuscript. Thanks also to Paul H. Arnaud, Jr., Mont A. Cazier, Joel F.
Gustafson, John S. Hensill, George E. Lindsay, and James R. Sweeney for
years of encouragement of my studies on scorpions.

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