545

ARTICLE
Importance of body size and hunting strategy during interactions
between the Mexican red-rump tarantula (Brachypelma vagans) and
the wolf spider Lycosa subfusca
A. Dor and Y. Hnaut
Can. J. Zool. Downloaded from www.nrcresearchpress.com by University of Toronto on 08/05/13

Abstract: Behavioural adaptation helps animals to maximize their ability to obtain food and to avoid being eaten, increasing
tness. To achieve this, they must assess predation risk and evaluate foraging needs simultaneously. In two sympatric spider
species, the wandering wolf spider Lycosa subfusca F.O.P. Cambridge, 1902 and the sit-and-wait Mexican red-rump tarantula
(Brachypelma vagans Ausserer, 1875), we studied the relationship between predatory behaviour and antipredatory behaviour at
different life stages. In the laboratory, encounters were organized between one wolf spider (small, medium-sized, or large) and
one tarantula (spiderling, small, medium-sized, or large). Attack latencies and behaviours were recorded. The results showed
that wolf spiders attacked and successfully captured younger tarantulas, while they avoided or retreated from older ones.
Tarantulas preferentially attacked and captured older wolf spiders. On other hand, younger wolf spiders were more cautious
than older ones, which waited until for the tarantulas to attack before retreating. Younger tarantulas were also more cautious
than adults, which never retreated from attack and increased their success in attacks with age. Finally, we discuss the relation-
ship between the predatory strategies of both spiders with their perception abilities and life history.

Key words: wolf spider, Lycosa subfusca, Mexican red-rump tarantula, Brachypelma vagans, Yucatan Peninsula.
For personal use only.

Rsum : Les adaptations comportementales des animaux optimisent lobtention de nourriture ou leur vitent dtre mangs,
en amliorant leur tness. Pour cela, ils doivent valuer simultanment le risque dtre mangs et la ncessit de se nourrir. Nous
avons tudi les comportements prdateur et anti-prdateur de deux araignes sympatriques, laraigne-loup errante Lycosa
subfusca F.O.P. Cambridge, 1902 et la mygale Brachypelma vagans Ausserer, 1875 pendant diffrentes priode de leur vie. En
laboratoire, on a ralis des rencontres entre une araigne-loup (petite, moyenne ou grande) et une mygale (trs petite, petite,
moyenne ou grande). On observe que les araignes-loups attaquent et capturent avec succs les plus jeunes mygales, alors
quelles nattaquent pas les adultes. Les mygales attaquent et capturent prfrentiellement les araignes-loups adultes. Dautre
part, les jeunes araignes-loups sont plus prudentes, elles vitent les mygales alors que leurs adultes attendent une attaque pour
se retirer. De mme, les jeunes mygales se montrent plus prudentes que leurs adultes, qui ne se retirent jamais en cas dattaque
et dont le succs dattaque augmente avec lge. Finalement nous discutons des relations entre les stratgies prdatrices de
chaque espce daraigne en lien avec leur habilet perceptive et leur histoire de vie.

Mots-cls : araigne-loup, Lycosa subfusca, mygale, Brachypelma vagans, pninsule de Yucatan.

Introduction
Predation is clearly one of the major selection pressures that
determine the form (Endler 1991), colour (Oxford and Gillespie
1998), and the behaviour of animals (Lima and Dill 1990; Lima
1998). Through the evolutionary arms race (Dawkins and Krebs
1979), predator and prey species maximize abilities to obtain food
or to avoid being eaten. This interaction has resulted in early
morphological adaptations in prey (Bengtson and Zhao 1992;
Bengtson and Hou 2001). Behavioural adaptations are also advan-
tageous against predators, because any animal that avoids preda-
tors or ees when it is attacked by predators has a greater
probability of surviving to mate and to produce offspring (i.e.,
tness) than other animals (Lind and Cresswell 2005). Neverthe-
less, excessive antipredatory behaviour is not necessarily synony-
mous with higher tness, as this type of behaviour or display is
generally costly because it interferes with foraging, mating,
and territorial defence (Lima and Dill 1990; Lima 1998; Lind and
Cresswell 2005). Optimizing the trade-off between foraging and
vigilance, prey should show graded responses of defensive behav-
iour proportional to the perceived risk (threat-sensitive predator
avoidance hypothesis; Dill and Fraser 1997; Sih 1997; Puttlitz
et al. 1999), and depending on a variety of intrinsic prey factors
such as development rate, feeding opportunities, mating oppor-
tunities, investment in offspring, body condition, and the risk
represented by the predator (Johnson and Sih 2007). If there is
little food available, prey is likely to take risks to forage in areas of
higher predation probability and is likely not to detect the attack-
ing predator. To resolve this dilemma, animals must assess preda-
tion risk and evaluate foraging needs concurrently, balancing the
two to maximize tness (Verdolin 2006).
Although many authors have focused their work on the anti-
predatory behavioural responses of prey against predators, few
studies are concerned with the relationship between the preda-
tory behaviour of a predator and its antipredator behaviour
against a superior predator. Indeed, in a system including a supe-
rior predator preying on an intermediate predator, the interme-
diate predator is simultaneously confronted with prey and a
superior predator. Its predatory strategies and antipredatory re-
sponses are shaped in a trade-off between preying and being
preyed upon (Schaffner and Anholt 1998). Because antipredatory

Received 3 December 2012. Accepted 9 May 2013.

A. Dor and Y. Hnaut. El Colegio de la Frontera Sur, Avenida Centenario Km 5.5, Chetumal 77014, Quintana Roo, Mxico.
Corresponding author: Ariane Dor (e-mail: ariane.dor@gmail.com, ador@ecosur.mx).

Can. J. Zool. 91: 545553 (2013) dx.doi.org/10.1139/cjz-2012-0308 Published at www.nrcresearchpress.com/cjz on 13 May 2013.
 546
and predatory strategies result from the evolutionary history of
predatorprey interactions (Lima and Dill 1990; Stapley 2004), we
may expect that the predatory strategy and the antipredatory
strategy are closely related to limit the cost that may result from
two distinct strategies.
The behavioural capacity of a predator to catch several types of
prey may allow it to have different behavioural responses to its
own predators. On the other hand, under the predation pressure
of a specic predator, a prey could have evolved to develop a
specic and efcient form of defence against this predator, but
inefcient with other predators (see Sih 1992). For instance, gen-
eralist predators such as burrowing tarantulas (family Therapho-
sidae) spend almost all their life in a protective burrow and shed
urticating setae from the abdomen by rapid downward strokes of
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the fourth pair of legs to repel predators (Cooke et al. 1972). These
antipredatory strategies seem to be efcient only against verte-
brate predators, which cannot enter the burrow and are repelled
by the urticating hairs. Wasps of the family of Pompilidae, a ta-
rantula parasitoid, can extract the spider from its burrow and
immobilize it (Punzo 1994) or attack and immobilize the spider in
the burrow (Costa et al. 2004). Defensive posture is another anti-
predatory behaviour that the tarantula uses against potential
predators, whether vertebrate or invertebrate (Costa et al. 2004;
Prez-Miles et al. 2005). In the event of a predator attack, like
many spiders, the tarantula can shed a limb that has been grasped
by the predator and thereby escape (Foelix 1996), the missing leg is
then regenerated on subsequent moults.
Wolf spiders (family Lycosidae) are wandering or sit-and-wait
generalist predators. They are the prey of vertebrates (lizards,
For personal use only.
birds, and small insectivorous mammals) and arthropods, in par-
ticular other spiders (Wise 1993, 2006; Foelix 1996; Dor and
Hnaut 2011). They have developed a range of defences against
predation and cannibalism based on chemical signals (Persons
et al. 2002; Wise 2006). We expect that they reduce the probability
of being detected by a predator by living in a burrow, as other
spiders do (Cloudsley-Thompson 1995). If a predator is close to
them, they are likely to avoid it before it is able to detect them
(Cloudsley-Thompson 1995; Persons et al. 2002) by reducing loco-
motion, increasing cover-seeking, or increasing vigilance (Lima
and Dill 1990; Lima 1998). When a predator nds them, they can
ee, running or jumping (Dor and Hnaut 2011), and when a pred-
ator attacks, they may shed the seized limb and escape (Foelix
1996).
Considering the problem of being a predator that may be
preyed upon by a superior predator, in terms of behavioural ad-
aptation costs, we focused our study on the predatory and anti-
predatory strategies of two spiders from the Yucatan Peninsula:
the burrowing Mexican red-rump tarantula (Brachypelma vagans
Ausserer, 1875) (Araneae: Theraphosidae) and the terrestrial wolf
spider Lycosa subfusca F.O.P. Cambridge, 1902 (Araneae: Lycosidae).
These spiders are generalist nocturnal terrestrial predators
(Marshall 1996; Toft and Wise 1999a, 1999b). The large adult taran-
tula B. vagans preys on L. subfusca (Dor and Weissenberger 2005;
Dor and Hnaut 2011), scorpions (Dor et al. 2011), ground-dwelling
arthropods (Machkour-MRabet et al. 2007), and small vertebrates
(Marshall 1996). The tarantula could be considered a superior
predator to the wolf spider, which is an intermediate predator.
The predatory strategies of the two spiders are distinct; tarantula
adults have a nightly sit-and-wait hunting strategy at the entrance
of the burrow (Machkour-MRabet et al. 2007), whereas adults of
L. subfusca, smaller and more active than tarantulas (Dor and
Hnaut 2011), engage in active hunting strategies in the eld dur-
ing the day and at night (Toft and Wise 1999a, 1999b). Antipreda-
tory strategies are not well known, but we expect tarantula adults
to display defence strategies against vertebrate predators through
its protective burrow and urticating abdominal hairs. We also
assume that adults of L. subfusca, active hunters, may be most
likely confronted with other organisms, as prey or predators, like
Can. J. Zool. Vol. 91, 2013
other wolf spiders including conspecics (Rypstra and Samu 2005;
Wise 2006; Dor and Hnaut 2011). It is known that wolf spider
species have developed adaptive antipredatory behaviour such as
prolonged periods of immobility, reduced walking speeds, and
avoidance of certain substrates (see Persons and Rypstra 2001;
Persons et al. 2001; Barnes et al. 2002).
Predatory and antipredatory strategies of early stages of both
spiders are little known. During the rst days of life, L. subfusca
spiderlings are all located on the top of their mothers back, with-
out any apparent aggressiveness toward their kin (A. Dor, per-
sonal observation), as many other spiders do (Gundermann et al.
1986; Jeanson et al. 2004). In natural conditions, the young
L. subfusca approach tarantula burrows less than older individuals
(see Dor and Hnaut 2011). In the case of B. vagans, before dispers-
ing, spiderlings live together in their mothers burrow, without
aggressiveness, although it seems that they eat cadavers of their
kin (Dor and Hnaut 2012). During dispersion from the maternal
burrow, a long column of about 100 tiny kin individuals (Shillington
and McEwen 2006), B. vagans spiderlings are particularly exposed
to predators. Once established in a burrow, preadult and adult
B. vagans are highly aggressive towards congeners (Hnaut and
Machkour-MRabet 2005; Dor et al. 2008).
The aim of the study is to demonstrate that two sympatric spe-
cies display different predatory and antipredatory strategies
along their life, and that shifts can be related to shifts in life
history: (1) adults of wolf spider, as wandering predators, have
conserved cautious behaviour of early stages and have developed
strategies based on decision making in respect to its own age and
its adversary, whereas (2) adults of B. vagans, as sit-and-wait pred-
ators, lost cautious behaviour of spiderlings, favouring attack on
all wolf spiders that approach without prior evaluation. We ven-
tured three hypothesis: (1) for both spider species, early stages are
more cautious and less aggressive than adult stages; (2) in sub-
adult and adult stages of wolf spiders, cautious behavior is con-
served but aggressiveness increases when they are larger than
tarantulas; and (3) in subadult and adult stages of tarantula, cau-
tious behaviour disappears and aggressiveness increases without
relation to their size ratio with wolf spider.
Materials and methods
Spiders maintenance
Tarantulas were collected in the village of Once de Mayo
(1810.9=N, 8945.9=W), on the eastern edge of the Calakmul Bio-
sphere Reserve (Campeche, Mexico), and wolf spiders were col-
lected on the ground around the compounds of El Colegio de la
Frontera Sur (1832=N, 8849=W) in Chetumal city (Quintana Roo,
Mxico). We reared them in the laboratory from several days to
several weeks between April 2005 and September 2006. The spi-
ders were maintained under the following laboratory conditions:
one individual per plastic box (13 cm in diameter and 5 cm high),
containing a cup lled with water to keep the humidity high. They
were fed with live larvae of superworms (Zophobas morio Fabricius,
1776) (Coleoptera: Tenebrionidae). Room temperature was main-
tained at 26 C, similar to natural conditions (Reichling 2003).
Prior to the experiments, the spiders were food-deprived for
1 week. The body size of each spider was determined by measuring
with a ruler the distance from the extreme anterior point of the
head to the hindmost part of the abdomen (spinnerets excluded).
Size classes were determined for each species to transform con-
tinuous variables into categorical variables (Table 1). We deter-
mined three size classes of wolf spider (small, medium-sized, and
large), and four size classes of tarantula (spiderling, small,
medium-sized, and large). Small wolf spiders presented a similar
size to tarantula spiderlings, but they were tinier than small ta-
rantulas. Medium-sized wolf spiders were equal in size with small
tarantulas, and smaller than medium-sized ones. Large wolf
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 Dor and Hnaut
Table 1. Size classes (mean SE) of the wolf spider Lycosa subfusca (three classes) and the Mexican red-rump tarantula (Brachypelma vagans) (four
classes) used in the experimental setup, with N being the number of spiders measured.
Mexican red-rump tarantula
Spiderling
(0.77 0.19 cm; N = 12)
Wolf spider
Small (0.91 0.20 cm; N = 20) NS
Medium-sized (43 0.28 cm; N = 20) ***
Large (2.08 0.07 cm; N = 20) ***
Note: Signicance tested with a MannWhitney U test. ***, P < 0.001; NS, not signicant.
spiders were bigger than small tarantulas and smaller than me-
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dium ones.
Observations of predatory and antipredatory behaviours
To test whether antipredatory and predatory behaviour of both
spiders are similarly affected by the size of the other spider,
we ran a series of encounters called trial under laboratory
conditions in which we confronted one wolf spider (small,
medium-sized, or large) with one tarantula (spiderling, small,
medium-sized, or large) and observed their behaviour.
All trials were performed at night (19000000) in clear plastic
boxes (29.5 cm wide 44 cm long 23.5 cm high) in which two
spiders were released gently and simultaneously at either end of
the box. All size combinations were tested: small wolf spider
(WS) spiderling tarantula (T), small WS small T, small WS
medium-sized T, small WS large T; medium-sized WS spider-
ling T, medium-sized WS small T, medium-sized WS medium-
For personal use only.
sized T, medium-sized WS large T; large WS spiderling T, large
WS small T, large WS medium-sized T, and large WS large T.
We then recorded the behaviour of both spiders using the fol-
lowing categories: approach, if one spider moved slowly toward
the other; avoidance, when one spider moved away from the
other before any attack occurred (attack did not happen); attack,
when one spider moved suddenly towards the other and made
contact with it; capture, when a spider bit another one after
attacking it; and retreat, when an attacked spider ran away,
without having been successfully grabbed or bitten. If they did not
move for more than 10 min, since the introduction of the spiders
into the box and they groomed themselves and stayed immobile,
we recorded these behavioural acts in a single category: keeping
quiet and the observation was considered nished. Those behav-
ioural acts were determined from preliminary observations with
both species and previous published works (Hnaut et al. 2001,
2005; Dor and Hnaut 2011; Dor et al. 2011) and are clearly distinct.
For each trial, we recorded the absence (0) or occurrence (1) of the
behaviour. Each trial lasted until either of the spiders retreated or
was captured. If neither of these two behaviours occurred, the
trial ended after 30 min of observations. Each of the 12 categories
of spider trial was repeated 30 times, except for trials involving
tarantula spiderlings, which were each repeated 10 times, because
of the lack of available spiderlings. Each trial involved a different
wolf spider; tarantulas, except for spiderling and small individu-
als, were removed from the arena as soon as a trial was over, and
were food-deprived again for a week. If it did not succeed to catch
and eat a wolf spider, a Z. morio larvae were put in its plastic box
and tarantula were kept quiet for a week. The time that elapsed
between the introduction of the individuals and the attack was
recorded. One observer recorded the behavioural data by direct
observations. Experiments were performed during the period
April 2005 to September 2006.
Data analysis
For each category of spider trial (for instance, small wolf
spider spiderling tarantula), we summed the occurrences for
each behaviour, obtaining behavioural frequencies in each cate-
gory of trials. To assess the hypothesis that early stages of wolf
547
Small Medium-sized Large
(1.32 0.35 cm; N = 14) (3.45 0.43 cm; N = 11) (4.53 0.32 cm; N = 15)
*** *** ***
NS *** ***
*** *** ***
spiders and tarantulas were cautious and little aggressive, we in-
dependently analysed behaviours of each spider as a function of
their size. For wolf spiders, we added together the behavioural
frequencies and attack latencies as a function of their size classes.
Therefore, we obtained three size groups: small, medium-sized,
and large wolf spiders. For tarantulas, we did the same, obtaining
four size groups of spiderling, small, medium-sized, and large
tarantulas. In the three size groups formed by wolf spiders and the
four size groups of the tarantulas, we also calculated an index
corresponding to attack success (number of capture / number of
attack). For analyses, we compared the behaviours of avoidance
and attack between wolf spider size groups with likelihood ratio
G test. We made one ow diagram for each wolf spider group. The
sum of percentages at the bottom of each diagram equals 100%.
Finally, we compared attack latencies with nonparametric
KruskalWallis tests followed by a Dunns test (Zar 1999), or
MannWhitney test. We did the same for tarantulas.
To assess the hypothesis that the wolf spider made the different
decisions as a function of the size of the tarantula, we added
together behavioural frequencies and attack latencies of the ta-
rantula as a function of their size classes, obtaining four size
groups of spiderling, small, medium-sized, and large tarantulas.
We compared the behaviours of avoidance and attack between
each size group of tarantula with a likelihood ratio G test. We
made one ow diagram for each tarantula group. Finally, we com-
pared their attack latencies with nonparametric tests.
To assess the hypothesis that the tarantulas increase their ag-
gressiveness without relation to their size ratio with wolf spiders,
we performed the same analyses as for wolf spiders, but in this
case, we control whether the tarantula behaviours depend on the
three wolf spider size groups. As tarantulas were at rest for a week
prior to any repetition and were randomly exposed to the differ-
ent spider size classes, we assumed that trials involving the same
individuals at a later date were independent. All tests were per-
formed using Statistica version 7 (StatSoft, Inc., Tulsa, Oklahoma,
USA) and values are reported as mean SE unless otherwise
reported.
Results
Cautious and less aggressive earlier spider stages?
Wolf spider
Small wolf spiders showed avoidance signicantly more than
attack (55% and 5%, respectively; Fig. 1A), as did medium-sized wolf
spiders (38% and 4%, respectively). Large wolf spiders avoided (21%)
and attacked tarantulas (17%) with similar frequencies.
Small, medium-sized, and large wolf spiders approached taran-
tulas with similar frequencies (Figs. 2A, 2B, 2C ). Small wolf spiders
avoided more confrontation with tarantulas than large wolf spi-
ders. Large wolf spiders attacked and captured more tarantulas
than small ones. Attack success was 60%, 100%, and 70% for small,
medium-sized, and large wolf spiders, respectively. Moreover,
medium-sized and large wolf spiders managed to capture few
tarantulas that attacked them. Wolf spiders practically never
retreated after they attacked a tarantula. Large wolf spiders
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 548
Fig. 1. (A) Percentage of avoidance (dark grey) and attack (light grey)
made by small, medium-sized, and large wolf spiders Lycosa subfusca
when confronted with Mexican red-rump tarantulas (Brachypelma
vagans) (regardless to their size). (B) Percentage of avoidance (dark
grey) and attack (light grey) made by spiderling, small, medium-
sized, and large tarantulas when confronted with wolf spiders
(regardless to their size). Behaviours were compared using a
likelihood ratio G test, where NA is a not applicable test and NS is a
not signicant test. ***, P < 0.001.
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For personal use only.
retreated more after an attack initiated by a tarantula than
smaller ones.
Small, medium-sized, and large wolf spiders had similar attack
latencies when confronted with tarantulas (9.3 2.0, 8.1 3.0, and
11.3 1.8 min; KruskalWallis test, H = 0.93, df = 2, n = 26, P = 0.63).
Tarantula
Tarantula spiderlings avoided wolf spiders (10%; Fig. 1B) as often
as they attacked (13%), whereas small tarantulas avoided wolf spi-
ders (2%) signicantly less than they attacked them (36%).
Medium-sized and large tarantulas never avoided confrontations
with wolf spiders and they attacked them in 51% and 53% of trials,
respectively.
Spiderlings, small, and medium-sized tarantulas approached
wolf spiders more than large tarantulas did (Figs. 3A, 3B, 3C, 3D).
Spiderlings and small tarantulas avoided wolf spiders to a similar
degree, while larger tarantulas did not avoid confrontations.
Larger tarantulas attacked and captured more wolf spiders than
smaller tarantulas. Attack success showed an increasing tendency
with tarantula size: 23% (spiderlings), 50% (small), 40% (medium-
sized), and 62% (large). Tarantulas never retreated from an attack
that they had initiated. However, when wolf spiders attacked,
smaller tarantulas tended to retreat.
Spiderlings, small, medium-sized, and large tarantulas attacked
with the same time intervals (3.8 0.5, 12.1 1.9, 10.3 1.3, and
9.2 1.4 min, respectively; KruskalWallis test, H = 4.407, df = 3, n =
110, P = 0.22).
Can. J. Zool. Vol. 91, 2013
Decision making as a function of the size of the adversary
Wolf spiders
When wolf spiders were in the presence of tarantula spider-
lings, they attacked them signicantly more than they avoided
them (57% and 13%, respectively; Fig. 4A). On the other hand, wolf
spiders avoided tarantulas signicantly more than they attacked
them when tarantulas were small (45% and 9%, respectively) and
medium-sized (39% and 1%, respectively). They never attacked
large tarantulas and avoided them in 39% of trials.
Wolf spiders approached the four size classes of tarantula sim-
ilarly (Figs. 3A, 3B, 3C, 3D). They avoided the spiderlings less than
small, medium-sized, and large tarantulas. They attacked and cap-
tured more spiderlings than larger tarantulas. Attack success was
higher on tarantula spiderlings than on small tarantulas (88% and
56%, respectively) and was null on medium-sized tarantulas. Wolf
spiders almost never retreated from their own attacks. They re-
treated more after tarantula attacks, although without relation to
tarantula size. Note that some wolf spiders managed to capture
small tarantulas in 2% of trials when those tarantulas attacked
them (Fig. 3B).
Wolf spiders tended (although not signicantly) to take longer
to attack spiderlings than small tarantulas (11.0 1.3 and 7.6
2.6 min, respectively; MannWhitney test, U = 36.5, df = 1, P =
0.067).
Tarantula
When tarantulas were in the presence of wolf spiders, they
always attacked them more than they avoided them regardless of
the wolf spiders size (Fig. 4B): small (29% attack and 3% avoid-
ance), medium-sized (47% attack and 1% avoidance) and large (55%
attack and 1% avoidance).
Tarantulas approached and avoided few wolf spiders and the
frequency of these behaviours was not inuenced by the wolf
spider size (Figs. 2A, 2B, 2C). Tarantulas attacked more medium-
sized and large wolf spiders than small wolf spiders, although
capture frequencies did not reect this tendency. Tarantulas
never retreated from their own attack and almost never retreated
from those initiated by wolf spiders. Tarantulas attack successes
on small, medium-sized, and large wolf spiders were similar (59%,
56%, and 40%, respectively).
Tarantulas attacked medium-sized wolf spiders more quickly
than large ones (11.2 1.3 min (small wolf spider), 7.2 1.3 min
(medium-sized wolf spider), 11.8 1.3 min (large wolf spider);
KruskalWallis test: H = 0.0114, n = 110, df = 2, P < 0.05).
Discussion
This study demonstrated that two sympatric spiders prey on
each other mutually depending on their size ratio, with smaller
individuals being killed by larger ones. Previous studies showed
that tarantulas regulated wolf spider natural populations through
predation of larger individuals (Dor and Hnaut 2011). In other
predatory species, intraguild predation also inuences popula-
tion structure and distribution (Hodge 1999; Wise and Chen 1999;
Suutari et al. 2004). Therefore, tarantula wolf spider mutual
predation may be considered a mutual stage-dependent intragu-
ild predation in which at least one of the two sympatric predators
regulates populations of the other predator (Polis and McCormick
1987; Polis et al. 1989; Polis 1991; Holt and Polis 1997). Others
factors which may affect predatorprey relationships is percep-
tion (Craig 1994; Heiling et al. 2005; Nakata 2010) and the behav-
ioural strategy (Craig 1994; Nakata 2007).
The wandering L. subfusca showed different behavioural strate-
gies with different sizes of B. vagans; they displayed adaptive be-
haviour modulated by their size ratio with the tarantula. The
ultimate aim of the adaptation seems to be the optimization of
their survival and (or) attack success, as they mainly focused their
attacks toward earlier stages of tarantulas. Young tarantulas are
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 Dor and Hnaut 549

Fig. 2. Flow diagrams of the behavioural sequences between three sizes of the wolf spider Lycosa subfusca (small (A), medium-sized (B),
large (C)) and the Mexican red-rump tarantula (Brachypelma vagans) (all sizes). For each label, the rst letters correspond with the spider,
i.e., tarantula (T) and wolf spider (WS). The behaviours that appear after the hyphen for each label correspond to the behaviours described in
the Materials and methods (Appr, approach; Atta, attack; Avoi, avoidance; Capt, capture; Retr, retreat; quiet). Broken arrows correspond to
percentage <10%, whereas solid arrows correspond to percentage <20% and boldface solid arrows correspond to percentage >20%. The sum of
percentages at the bottom of each diagram equals 100%.
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For personal use only.

as small as small wolf spiders and can be easily preyed on. More-
over, when wolf spiders attacked tarantulas, they captured them
in more than half the attacks. These attack successes are similar to
those previously observed in cannibal confrontations of L. subfusca
with a range from 50% to 80% (Dor and Hnaut 2011). When the
wolf spiders were smaller than the tarantulas, they were exposed
to the danger of a potential attack by tarantulas. Thus, young wolf
spiders chose to avoid confrontation, while older ones did not;
they rather retreated successfully from tarantula attack. These
observations are similar to results obtained in eld studies in
which the smaller L. subfusca curled up, prostrated, or avoided
confrontation with burrowed B. vagans, while larger wolf spiders
retreated from more than 70% of the tarantulas attack (Dor and
Hnaut 2011). In standard conditions, more confrontations oc-
curred than in the eld, undoubtedly because it was impossible
for the wolf spider to use hiding places in its environment that
made them unnoticeable to the sit-and-wait tarantula as is the
case for other organisms (Lensing and Wise 2004; Rosenheim
2005; Finke and Denno 2006). Older wolf spiders exceptionally
managed to capture small tarantulas when the latter attacked

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 550 Can. J. Zool. Vol. 91, 2013

Fig. 3. Flow diagrams of the behavioural sequences between four sizes of the Mexican red-rump tarantula (Brachypelma vagans) (spiderling (A),
small (B), medium-sized (C), large (D)) and the wolf spider Lycosa subfusca (all sizes). For each label, the rst letters correspond to the spider, i.e.,
tarantula (T) and wolf spider (WS). The behaviours that appear after the hyphen for each label correspond to the behaviours described in
the Material and methods (Appr, approach; Atta, attack; Avoi, avoidance; Capt, capture; Retr, retreat; quiet). Broken arrows correspond to
percentage <10%, whereas solid arrows correspond to percentage <20% and boldface solid arrows correspond to percentage >20%. The sum of
percentages at the bottom of each diagram equals 100%.
Can. J. Zool. Downloaded from www.nrcresearchpress.com by University of Toronto on 08/05/13
For personal use only.

them. It is comparable with the situation when tarantulas were
attacked by yellow and black scorpions; B. vagans also managed to
capture them in some trials (Dor et al. 2011). In both cases, success-
ful tarantulas and wolf spiders were larger than their respective
enemies. Therefore, in our standard conditions, we demonstrated
that L. subfusca made decisions about the choice of the best behav-
iour to handle the tarantula depending on the size ratio of the two
animals. This decision is generally successful, because wolf spi-
ders died in a few cases.
The strategy of tarantulas was less related to the spiders sizes
and thus to decision making. Indeed, prey size is not always a
limiting factor. They did not approach wolf spiders, as they are
sit-and-wait hunters (Machkour-MRabet et al. 2007). The smallest
tarantulas avoided wolf spiders that were larger than themselves,
but when they attacked wolf spiders, they had little success. On
the contrary, the larger tarantulas attacked wolf spiders, but they
only managed to capture less than the half of the attacked spiders.
These attack successes were lower than those obtained in our

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 Dor and Hnaut
Fig. 4. (A) Percentage of avoidance (dark grey) and attack (light
grey) made by the wolf spiders Lycosa subfusca when confronted with
four sizes of Mexican red-rump tarantulas (Brachypelma vagans)
(spiderling, small, medium-sized, and large). (B) Percentage of
avoidance (dark grey) and attack (light grey) made by the tarantulas
when confronted with three sizes of wolf spiders (small, medium-
sized, and large). Behaviours were compared using a likelihood ratio
G test, where, NA is a not applicable test. **, P < 0.01; ***, P < 0.001.
Can. J. Zool. Downloaded from www.nrcresearchpress.com by University of Toronto on 08/05/13
For personal use only.
previous study (Dor et al. 2011) for tarantulas attacking yellow and
black scorpions, crickets, and cockroaches. This difference in at-
tack success was related to the retreat acts of large wolf spiders
compared with other potential prey of B. vagans, such as offensive
scorpions. Indeed tarantulas, which preferentially attacked larger
wolf spiders, captured the same proportion of wolf spiders what-
ever their size. Another not exclusive explanation of the poor
efciency of tarantula attacks is that the tarantula adults can
withstand a period without any food of about 2 years (Baerg 1958),
whereas food-deprived wolf spider adults survived less than a year
(Anderson 1974). Moreover B. vagans may attack all potential prey
that they detect in their proximity. Machkour-MRabet et al. (2007)
found remains of ants, beetles, and hemipterans in B. vagans bur-
rows, while Marshall (1996) found small vertebrates. Finally, small
B. vagans seem to be more cautious than adults. This tarantula
does not appear to have developed decision-making behaviour to
catch prey as the wolf spider does. In fact, tarantula behaviour
looks more like a reex behaviour. The lack of prey evaluation
between detection and attack may explain why tarantulas are
heavily subject to parasitic wasps that takes advantage of this
automatic attack when vibrations are detected (Libersat et al.
2009).
The preference of tarantulas for larger wolf spiders can be ex-
plained by their limited detection abilities. Previous works dem-
onstrated the poor vibratory detection of prey by the Brazilian
salmon pink bird-eating tarantula (Lasiodora parahybana (Mello-
Leito, 1917)) (Redondo 1994; Blein et al. 1996) and its limited visual
abilities (Baerg 1958). For instance, the vibrations created by small
prey do not cause attack of the golden orb-web spider (Nephila
clavipes (L., 1767)) (Zschokke et al. 2006; Hnaut et al. 2010). Thus,
because the tarantula uses vibration to detect prey (Blein et al.
551
1996), it is probable that the vibrations created by the slow small
wolf spiders were not detected by the tarantula, whereas the con-
stantly moving cockroaches were easily detected (Dor and Hnaut
2011). A second reason could be that the small wolf spiders provide
fewer nutrients than larger cockroaches when predated. Spiders
have feeding preferences; they usually only catch prey that is 50%
80% of their own size; smaller prey items are typically ignored
(Nentwig and Wissel 1986; Nyffeler et al. 1994; Marc and Canard
1997; Marc et al. 1999; Hnaut et al. 2001, 2005; Zschokke et al.
2006). A third explanation maybe that tarantulas focus their at-
tention on limited foraging areas near its burrow entrance, as the
orb-web spider Cyclosa octotuberculata Karsch, 1879 does (Nakata
2010).
In the two species that we studied, the comparison of attack
success between two similar-sized individuals (i.e., small tarantu-
las and medium-sized wolf spiders) shows that wolf spiders were
more successful than tarantulas. Moreover, adult tarantulas pre-
sented higher attack success than juveniles. Therefore, small ta-
rantulas perhaps do not have the experience to catch prey such as
the wolf spider and express more cautious behaviour. So, size and
behaviour are not the only factors that inuence the interaction
issue; maturity level and (or) previous experience (Hnaut et al.
1999, 2000) and maturity level (Hendrichs et al. 2007) also do.
Another question arises concerning the inuence of the tarantu-
las maturity on its perception and thus on the decisions that they
make in the presence of a predator.
In addition, because wolf spiders wander and lie in wait in the
open eld to obtain food, they are constantly exposed to other
organisms, whether prey or predator. Although they display a
cryptic dark brown pattern on the cuticle (which helps them to
remain unnoticed in leaves or a dusty environment), they do not
have physical protection such as a burrow, or an offensive sting or
strong carapace as scorpions do, and their adult size is relatively
small compared with those of tarantulas. Therefore, decision
making and adaptive behaviour in the presence of tarantulas are
key factors in their predatory and antipredatory strategies. They
have developed complex behavioural responses to the presence of
a tarantula based on the perception of the size, or the potential
danger the other organism represents. This behaviour seems to be
innate because small wolf spiders did it; however, retreat behav-
iour may be a learned behaviour based on experience (Nakata
2007). The greater capacity to retreat may also be a result of age
and the development of the antipredatory abilities (Jedrzejewska
and Jedrzejewski 1990; Hendrichs et al. 2007).
On the other hand, tarantulas show the opposite tendency.
Large tarantulas count on a protective perennial burrow, they
release urticating hairs when they feel threatened, and they at-
tack larger preys. They never express cautious behaviour such as
avoidance or retreat except during their early stages, so they can
fall prey to wolf spiders. This shift of behaviour between early
stages and adulthood can be related to the shift from a young
wandering to an adult sedentary lifestyle (Fuiman and Magurran
1994). Spiderling tarantulas disperse from the maternal burrow
along conspicuous columns searching for a new shelter (Reichling
2000; Shillington and McEwen 2006; Dor and Hnaut 2012). Small
tarantulas are often encountered in the eld or in supercial
cavities (A. Dor, personal observation). Consequently, they are
exposed to potential predators such as wolf spiders and this could
be related to their cautious behaviour. In conclusion concerning
the shift of behaviour in tarantulas, behaviour seems to have a
variable role depending on the maturity stage and hence the life-
style, and essentially depends on their perception abilities.
Therefore, as large tarantulas do not perceive smaller wolf spi-
ders, it seems that regulation of wolf spider populations by the
tarantula (Dor and Hnaut 2011) is perception dependent. It regu-
lates only larger wolf spider individuals, limiting their reproduc-
tive potential and the recruitment of juveniles. The predation by
wolf spiders on early stages of tarantula acts differentially on
Published by NRC Research Press
 552
tarantula populations, as it limits the dispersion efciency of
B. vagans, explaining partially why tarantula populations have a
patchy distribution (Machkour-MRabet et al. 2005).
In conclusion, two sympatric spiders, very similar in appear-
ance, adopt different strategies depending on their adult sizes and
modes of life. Large B. vagans adults are generally positioned at the
entrance to their burrow, from where they attack prey by biting.
In contrast, L. subfusca and spiderlings of B. vagans rst have to
detect the approaching potential predator to avoid it eeing, curl-
ing up, or hiding in available crevasses in the eld. Consequently,
living in a burrow seems to diminish the necessity for decision
making in B. vagans, whereas living in the open ground, L. subfusca
is constantly under predation pressure and must make decisions
to insure its survival. A remaining question is whether the bur-
Can. J. Zool. Downloaded from www.nrcresearchpress.com by University of Toronto on 08/05/13
rowing life has caused the loss of cognitive capacities of B. vagans.
The fact that adult females of this species are found wandering
around and are able to attack and kill burrowing congeners to
steal their burrow (Hnaut and Machkour-MRabet 2005) provides
an element of response to this question. In the open environment,
female B. vagans are chemically attracted towards burrowing fe-
male congeners; this attraction seems to confer wandering fe-
males an advantage over burrowing ones (Dor et al. 2008). Thus,
B. vagans does not lose cognitive abilities; instead it adapts its
behavioural responses as a function of its environment. On the
other hand, wolf spiders generally seem to have developed anti-
predatory strategies based on perception of dangerous organisms
through vision, vibration sensitivity (Rovner 1989, 1996), and
chemical perception (see Persons and Rypstra 2001). Therefore, we
could put forward the hypothesis that L. subfusca perceives
For personal use only.
B. vagans (for instance, it could see the B. vagans shape, perceive its
vibrations, and (or) its kairomones), obtaining information about
its size and its danger level. Depending on the information per-
ceived, the wolf spider will make a decision about the more ap-
propriate tactic: avoidance or attack. Experience can modulate
their behaviour between the choice of avoidance or retreat.
It is also possible that the wolf spider makes a search image
(Tinbergen 1960) of a particular type of prey that it will prey upon,
like the jumping spider Portia labiata (Thorell, 1887) (Jackson and Li
2004). We can wonder if wolf spiders are able to make a search
image of B. vagans spiderlings and which sensory canal(s) is used
for. Moreover, it would be interesting to determine the variable
characteristic(s) of B. vagans that modify(ies) the wolf spider behav-
iour from attack to avoid and how it is perceived by the wolf
spider. If such a search image exists for the wolf spider, then what
would be the threshold to switch between searched prey (corre-
sponding to the search image) and avoided predator (not corre-
sponding to the search image) of the wolf spider?
Acknowledgements
This work was only possible with the technical help of D.
Lecocq, N. Snchez, J. Perez Campos, and L. del Roco, and the
friendly collaboration of the people of the village Once de Mayo. L.
Legal, P. Winterton, C. Shillington, and two anonymous reviewers
are specially thanked for valuable comments on an earlier version
of the manuscript. The experiments complied with the current
laws of the country in which they were performed. Financial sup-
port was provided by the Mexican Consejo Nacional de Ciencia y
Tecnologa (CONACYT, Project H-52113).
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