ethology
international journal of behavioural biology
A Continuum of Behavioral Plasticity in Urban and Desert Black
Widows
Rebecca N. Halpin & J. Chadwick Johnson
School of Mathematical and Natural Sciences (2352), Arizona State University at the West Campus, 4701 W. Thunderbird Road, Glendale, AZ 85306,
USA
Correspondence
Rebecca N. Halpin, 16185 West Lincoln
Street, Goodyear, AZ 85338, USA.
E-mail: Rebecca.n.halpin@gmail.com
Received: June 3, 2014
Initial acceptance: July 7, 2014
Final acceptance: July 17, 2014
(S. Foster)
doi: 10.1111/eth.12297
Keywords: behavioral plasticity, behavioral
syndromes, human-induced rapid
environmental change, Latrodectus hesperus
Introduction
Recently, behavioral ecology has experienced a resur-
gent focus on the importance of the behavioral plas-
ticity expressed across an organism’s behavioral
repertoire (e.g., Gross et al. 2010; Miranda et al.
2013; Salinas-Melgoza et al. 2013) which marks the
ability of an organism to modulate their behavior such
that they respond adaptively within distinct contexts
(Dingemanse et al. 2009). This resurgence is
particularly important in the context of ecological dis-
turbance caused by human-induced rapid environ-
mental change (HIREC). HIREC presents the ‘grand
challenge’ of understanding how individual organ-
isms cope with human-induced changes (Vitousek
et al. 1997; Sih et al. 2010b; Sih 2013). For example,
Ethology 120 (2014) 1237–1247 © 2014 Blackwell Verlag GmbH
Ethology
Abstract
Behavioral plasticity marks an individual’s ability to modulate behavior
across functional contexts. Behavioral syndromes, on the other hand,
appear as consistent individual variation in behavior that is both repeat-
able for individuals within a functional context (e.g., consistent voracity
toward prey) and correlated across contexts (e.g., high voracity toward
prey and high levels of boldness toward enemies). Thus, adaptive plastic-
ity and syndromes represent two extremes of a behavioral plasticity con-
tinuum upon which most behavioral phenotypes fall. We tested for both
adaptive plasticity and behavioral syndromes in the western black widow
spider, Latrodectus hesperus. We measured behavior in three contexts: star-
tle, startle + prey, and startle + mate, and found (1) classic behaviorally
plastic responses to predation risk, (2) high repeatability of behavior
within contexts, and (3) evidence of a correlation between startle + prey
and startle + mate contexts, indicative of a behavioral syndrome. As rela-
tive behavioral plasticity may vary across populations, we also compared
urban and desert populations to test whether spiders from these habitats
exhibit different behaviors and/or behavioral syndromes. While we found
that urban males used in mating trials courted urban females significantly
more than desert females, we found no other differences in the behavior
of urban and desert black widows. Thus, black widows, regardless of habi-
tat, are characterized by both context-specific behavioral plasticity and
across-context correlations, presenting a phenotypic complexity that is
likely exhibited, to varying degrees, by most organisms.
urbanization offers a prime example of HIREC, and
studying behavioral plasticity in urban-disturbed
environments (e.g., Miranda et al. 2013) can help us
understand how and why ‘urban exploiters’ (taxa that
exploit the urban environment and reach their great-
est population densities in urban areas; Blair 1996)
thrive during and after human disturbance.
In contrast, some organisms, display low levels of
behavioral plasticity. As such, a great deal of recent
work has focused on two areas of behavioral consis-
tency (Dall et al. 2004; Sih et al. 2004a,b; Bell 2007).
First, individuals can exhibit repeatability within a
behavioral context (Boake 1989). For example, an
individual that is highly voracious toward prey only
when it is in poor condition would show lower levels
of repeatability than another individual who
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Relative Plasticity in Black Widow Spiders
expresses high levels of voracity toward prey regard-
less of its feeding condition (see Bell et al. 2009 for
meta-analysis of behavioral repeatability). Repeatabil-
ity is important as it implies that either selection has
favored an unwavering phenotype or that there are
constraints placed on individuals that force them to
act similarly from one encounter with a stimulus to
the next.
Second, studies of behavioral consistency are also
concerned with whether behaviors are variable (i.e.,
plastic) or invariable (i.e., correlated) across different
functional contexts. When individual variation is both
repeatable within a context and correlated across
functional contexts, these suites of correlated behav-
iors have been termed ‘animal personalities’ (Gosling
1998; Dingemanse & Reale 2005; Bell 2007) or
‘behavioral syndromes’ (Sih et al. 2004a,b, 2010a).
For example, individual funnel web spiders, Agelenop-
sis aperta, show consistency in their level of aggression
across foraging, territoriality, and antipredator con-
texts (Riechert & Hedrick 1993), such that the most
voracious predators are also the most territorial
toward conspecifics and the boldest toward predators.
Similarly, a more social spider, Anelosimus studiosus,
demonstrates a behavioral correlation such that the
most agonistic (least social) spiders are also the most
aggressive toward prey and boldest toward predators
(Pruitt et al. 2010; see also Johnson & Sih 2005, 2007
for a similar example that includes sexual cannibalism
as a part of the behavioral syndrome).
If the correlations across contexts that constitute a
behavioral syndrome are shown to yield fitness bene-
fits, then a behavioral syndrome can be considered
adaptive (e.g., showing decreased boldness in the
presence of predators and being non-aggressive
toward conspecifics that participate in group antipre-
dator behaviors; Bell & Sih 2007). However, behav-
ioral syndromes may also be non-adaptive as
behavioral correlations across contexts may prevent
an individual from modulating their behavior and
behaving optimally in all contexts. Such correlations
could explain the existence of ‘inappropriate’ behav-
iors, such as high activity levels in the presence of pre-
dators (Sih et al. 2003) and the extreme sexual
cannibalism of all potential mates by female fishing
spiders (Arnqvist & Henriksson 1997; Johnson & Sih
2005).
While most studies of behavioral syndromes focus
on individual variation within a population, syn-
dromes can also vary across distinct populations
(Royaute et al. 2013). Such variation among popula-
tions can be caused by variable selective environ-
ments, mutation, genetic drift, founder effects, and
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R. N. Halpin & J. C. Johnson
gene flow (Dingemanse et al. 2007). For example,
Bell & Sih (2007) showed that stickleback populations
(Gasterosteus aculeatus) from high-risk environments
demonstrate strong behavioral correlations while
stickleback populations from low-risk environments
show a much higher degree of behavioral plasticity.
Thus, rigid behavioral syndromes may be most preva-
lent when they are the adaptive result of strong,
unwavering selection pressures (i.e., behavioral corre-
lations may only persist when ‘mistakes’ are costly).
In contrast, plastic phenotypes may be most com-
monly found where a species encounters weaker or
variable selection pressures (Sih 2013).
Urbanization offers an opportunity to examine
unique urban selection pressures and resultant popu-
lation behavioral differences between disturbed-
urban and undisturbed-rural populations (Miranda
et al. 2013). These differences in selection pressures
across urbanized and relatively undisturbed environ-
ments have been shown to have profound effects on
where a species falls along the behavioral plasticity
continuum. For example, Song Sparrows, Melospiza
melodia, from rural populations display behavioral
correlations between aggression and boldness behav-
iors, while those from urban populations show no evi-
dence of a behavioral syndrome (Evans et al. 2010;
Scales et al. 2011). This suggests that selection pres-
sures in urban habitats favor greater plasticity in this
species, resulting in the uncoupling of aggression and
boldness traits.
Spider taxa have frequently been used as a model
in studying behavioral syndromes (reviewed in Pru-
itt & Riechert 2012). Many spiders show high levels
of aggression, a trait that can be studied in a num-
ber of different functional contexts (e.g., antipreda-
tor, foraging, and mating). Furthermore, spiders
show ecotypic variation or differences due to local
adaptation, resulting in behavioral variation across
large geographic ranges (Riechert et al. 2001), thus
allowing for the study of behavioral syndrome dif-
ferences across ecologically distinct populations. The
western black widow spider, Latrodectus hesperus is
one spider system that allows for an examination of
behavioral responses to urban disturbance. Latrodec-
tus hesperus is an urban pest of medical importance
that infests urban habitats throughout the desert
Southwest (Trubl et al. 2012). The black widow’s
behavioral repertoire is particularly well suited to
the study of behavioral syndromes as they exhibit
high levels of aggression toward (1) predators (Vet-
ter 1980), (2) conspecifics (Johnson et al. 2010), (3)
prey (Trubl et al. 2010), and (4) mates (Johnson
et al. 2011).
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R. N. Halpin & J. C. Johnson
Here, we test for the presence of both traditional
adaptive behavioral plasticity and a behavioral syn-
drome in L. hesperus, across three functional contexts:
(1) startle response, (2) startle + prey response, and
(3) startle + mate response. Regarding adaptive plas-
ticity, we predict that individuals should be more
active in startle + prey and startle + mate contexts
relative to the startle context. In contrast, as per the
behavioral syndromes hypothesis, we predict that (1)
an individual’s behavior will be consistent (repeat-
able) across replicate measures within each of these
contexts and (2) behavioral correlations across these
three contexts will expose a behavioral syndrome. For
example, spiders that are most active after being star-
tled are predicted to be the same spiders that are the
most active when presented with prey and the most
active in the presence of potential mates. Addition-
ally, we compare these behaviors across urban and
desert populations predicting population differences
in both plasticity and behavioral syndromes. Relative
to arthropods from urban habitats, desert arthropods
are thought to experience stronger, more intense
selection pressures (Shochat et al. 2006). Therefore,
we predicted that desert spiders would display tight
behavioral correlations. In addition, because we note
that urban populations are typified by great spatial
variation (Trubl et al. 2010) and selection pressures
may be relaxed in urban areas, we predicted that
urban spiders would display a more plastic phenotype
than their desert counterparts.
Methods
We followed 20 adult, female black widows over the
course of 9 wk across three behavioral contexts: (1)
startle response (six repeated measures), (2) star-
tle + prey (four repeated measures), and (3) star-
tle + mate (four repeated measures). Ten females
were collected from urban sites across Phoenix, AZ.
The remaining ten spiders were collected from undis-
turbed Sonoran desert habitats more than 65 km
outside of Phoenix. Spiders were collected during the
last week of June 2012 and brought back to the lab-
oratory where they were weighed to the nearest mil-
ligram and fed two house crickets (Acheta domesticus),
each prey item being approx. 50% of the mass of the
spider. One week later, spiders were weighed again
and placed in identical laboratory environments,
designed to be unique when compared to both desert
and urban environments (Fig. 1). This was accom-
plished by using a 57 9 38 9 33 cm plastic tubs
with 1 cm of sand coating the tub’s floor. A
5 9 19.5 9 15.5 cm Styrofoam wall was glued to
Ethology 120 (2014) 1237–1247 © 2014 Blackwell Verlag GmbH
Relative Plasticity in Black Widow Spiders
Fig. 1: 57 9 38 9 33 cm laboratory environments with 1 cm of sand,
5 9 19.5 9 15.5 cm Styrofoam wall, 4.5 9 5 9 5.5 cm refuge, and
53 cm wooden dowels.
one end of the tub, and a refuge was created by
inserting a halved Styrofoam cup into a
4.5 9 5 9 5.5 cm cutout at the top of the Styrofoam
wall (this refuge structure allowed us to control for
both the position and approximate size of each spi-
der’s refuge as all spiders built their refuge within
the provided cup). Two 53 cm wooden dowels
extended the length of the tub from the refuge
entrance to the floor of the tub’s opposite end, acting
as scaffolding for web building. These dowels not
only assisted spider’s in web building within their
laboratory environments, but also allowed us to con-
trol for both the approximate size and shape of each
spider’s web as they were confined to building web
where scaffolding was provided (spiders could not
effectively attach web to the sides of the tub). Spi-
ders were allowed to construct their webs for 1 wk
before entering into a fixed sequence of repeated
behavioral trials (see below) that were identical for
each spider. All spiders were hand-fed one cricket
each week for the duration of the trials by holding
the cricket on the web until feeding occurred. Spi-
ders were weighed 24 h after each feeding. Behav-
ioral trials were always separated from one another
and from feedings by at least 48 h. Over the course
of our 3-mo trial period, 15 of 20 females produced
egg sacs (six urban, nine desert), indicating that
most, if not all, of the spiders in our study were col-
lected as mated adults.
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Relative Plasticity in Black Widow Spiders
Startle Response trials
Spiders underwent six 3-h startle response trials over
the course of 6 wk, allowing for 6 d between trials.
Startle response trials always began with the spider
being firmly prodded on the abdomen repeatedly with
a wooden dowel until they retreated to the back of
their refuge and assumed a defensive posture (legs
curled close to the abdomen). Individuals were moni-
tored every 2 min for the first hour and every 10 min
for the second and third hours. This allowed for the
calculation of the following measures of boldness:
proportion of trial females spent active (spiders were
considered active if they were moving at the time of
the spot check or if they had changed location from
the last spot check), latency to leave refuge posture
(time taken to transition from defensive/refuge pos-
ture in which legs are tightly curled around abdomen
into an open, foraging posture with legs extended into
the web), and average female distance from refuge.
Startle + Prey Trials
Females underwent four 3-h startle + prey trials over
the course of 4 wk, allowing for 6 d between trials. In
preparation for trials, individual crickets (Acheta do-
mesticus) were adhered to a 46 cm piece of thread by
hot gluing their pronotum to the end of the thread.
As described in the startle response trial procedure,
females were then disturbed until they retreated to
the back of the refuge. By disturbing each spider in
the same way, we were able to standardize the spi-
der’s location prior to each trial. Immediately after
this disturbance, crickets were dropped by their
thread into the tub at a distance of 30 cm from the
refuge. The opposite end of the thread was then taped
to the outside of each tub such that crickets were free
to move in the web but prevented from ranging
widely in the spider’s tub. These startle + prey trials
were scored every 2 min for the first hour and every
10 min for the second and third hour to measure the
percentage of spider activity, latency to leave refuge
posture, average distance from refuge, and latency to
contact prey. To prevent consumption of crickets, tri-
als were ended, and crickets were removed after spi-
ders had wrapped their prey for ten consecutive
minutes.
Startle + Mate Trials
Females underwent four 3-h startle + mate trials over
the course of 4 wk, allowing for 6 d between trials. A
pool of 45 adult males, each from different families,
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R. N. Halpin & J. C. Johnson
reared in the laboratory from parents collected from
urban populations was used to randomly pair a single
male with each of the 20 focal females. Males were
used in multiple trials throughout the course of the
experiment, but no male was ever paired with the
same female twice. As described for the startle
response trials, females were ‘disturbed’ until they
retreated to the back of their refuge. By disturbing
each spider in the same way, we were able to stan-
dardize the spider’s location prior to each trial. Imme-
diately after this disturbance, an adult male was
placed in the web 30 cm from the refuge. Mating tri-
als were scored every 2 min for the first hour and
every 10 min for the second and third hour to mea-
sure: percentage of trial spent active (male and
female), latency to leave refuge posture (female), and
distance from refuge (male and female). To prevent
an actual copulation, trials in which a male courted
on the female’s abdomen for 30 consecutive minutes
were ended and the male was removed (in this species
it is extraordinarily uncommon to see pedipalp inser-
tion occur within the first 30 min of abdomen court-
ship; J. C. Johnson, unpubl. data). In all but five of
the 80 startle + mate trials we conducted, males
began courting within 15 min of placement on the
female’s web. No females encountered more than one
male who failed to court within 15 min. Previous
work has shown that male courtship levels increase
significantly as female activity and aggression
decreases (Johnson et al. 2010). Thus, we recorded
male courtship activity throughout these trials as an
indirect measure of female courtship receptivity.
Statistical Analysis
To address our questions about potential behavioral
differences between urban and desert spiders, ANOVA
was used with habitat origin as an independent vari-
able. Subsequent analyses either pooled all 20 spiders
across habitat types if we found no differences across
habitats, or were performed separately for each habi-
tat type if significant differences between habitats
were found. A chi-square test was utilized to deter-
mine whether the number of egg sacs produced by
urban and desert populations significantly differed,
while a series of paired contrasts determined whether
there were significant behavioral differences between
spiders that produced egg sacs and those that did not.
MANOVA was utilized in an attempt to test for overall
significance between behavioral contexts. This
allowed for the use of multiple paired contrasts to test
for behavioral differences between individual behav-
ioral contexts without a Bonferroni adjustment.
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R. N. Halpin & J. C. Johnson Relative Plasticity in Black Widow Spiders

Intraclass correlation coefficients were used to test for

individual repeatability across repeated measures
within each behavioral context (Boake 1989) and a
Fisher’s r-to-z transformation (Zar 1984) was used to
determine the relative behavioral repeatability
between contexts (Sinn & Moltschaniwskyj 2005). To
test for the presence of behavioral correlations across
contexts, as predicted by the behavioral syndromes
hypothesis, a multivariate regression was first used to
test for overall correlations between all contexts. This
allowed for the use of multiple bivariate correlations to
test for the behavioral correlations across specific con-
texts without the use of a Bonferroni adjustment. Two-
tailed p-values were used in all statistical analyses.

Results
Fig. 2: Male spiders (descended from urban parents) spend more time
Desert and Urban Differences courting urban females than desert females (F1,18 = 4.782, p = 0.042).

Urban and desert female spiders showed no significant

differences in their activity levels, average distance Table 2: Behavioral differences between spiders that produced egg
from refuge, latency to leave refuge posture, and sacs and those that did not
latency to prey contact (see Table 1). Interestingly,
Measure Context t p
laboratory-reared male spiders used in our mating tri-
als (descended from urban parents) spent more time Percent activity Startle 0.195 0.424
courting urban females than desert females Startle + Prey 0.79 0.22
(F1,18 = 4.782, p = 0.042: Fig. 2). Hence, below, we Startle + Mate 0.76 0.228
Average distance from refuge Startle 0.587 0.282
analyze this measure separately for urban and desert
Startle + Prey 0.081 0.468
spiders. For all other analyses, we pooled urban and Startle + Mate 0.283 0.39
desert spiders into the same analysis. Latency to leave refuge posture Startle 0.039 0.485
We found no significant difference between the Startle + Prey 0.258 0.399
number of egg sacs produced by urban and desert spi- Startle + Mate 0.729 0.238
ders (v2ð1;N¼19Þ = 0.6, p = 0.439). Additionally, no sig- Number of harassments Startle 0.173 0.432
nificant behavioral differences were found between
spiders that produced egg sacs and those that did not tle + mate contexts (F2,17 = 15.73, p < 0.05). Specifi-
(see Table 2). cally, females were significantly more active in
startle + prey and startle + mate trials than in startle
Behavioral Plasticity Across Contexts response trials (startle + prey vs. startle response:
t = 5.16, p < 0.01; startle + mate vs. startle response:
We found significant differences across behavioral t = 4.91, p < 0.01), with no significant difference
contexts (F1,19 = 510.8, p = 0.035). For example, between startle + prey and startle + mate trials
female activity levels (Fig. 3A) varied significantly (t = 0.62, p = 0.62). We also found within-context
across startle response, startle + prey, and star- correlations that suggest female activity levels have
Table 1: Desert and urban site comparisons important ecological implications: individuals that
showed high levels of activity while foraging tended
Measure F1,18 p
to capture prey faster than those that exhibited lower
Percentage of female activity 0.277 0.605 levels of activity (r = 0.594, N = 20, p < 0.05), and
Latency to leave refuge posture 0.001 0.972 urban females that were more active in the presence
Average female distance from refuge 0.283 0.601 of a male tended to be courted less persistently by
Latency to prey contact 0.026 0.873 males (desert and urban spiders were separated for
Percentage of male courtship activity 4.782 0.042
this analysis to account for desert/urban behavioral
Number of harassments 0.008 0.928
differences discussed above; r = 0.805, N = 10,
Bold values are statistically significant. p = 0.005, Fig. 4).

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Relative Plasticity in Black Widow Spiders
(a)
(b)
Fig. 3: (A) Average differences across behavioral contexts in the per-
centage of time females spent active. (B) Average differences across
behavioral contexts in a female’s average distance from her refuge.
Fig. 4: Male courtship of active, urban females.
Our measure of a female’s average location in the
web (distance from her refuge) showed that females
resided at significantly greater distances from the
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R. N. Halpin & J. C. Johnson
refuge in startle + prey trials than either other trial
type (startle + prey vs. startle response: t = 3.93,
p < 0.01, startle + prey vs. startle + mate: t19 = 3.38,
p < 0.01; see Fig. 3B). No significant difference was
found between female average distance from refuge
in startle response and startle + mate trials (t = 0.34,
p = 0.37).
Behavioral Consistency (Repeatability) Within
Contexts
Regarding within-context repeatability, we found a
great deal of consistency within repeated measures of
the same trial type (Table 3). Our results showed that
the number of harassments required before a spider
retreated to its refuge was highly repeatable in the 14
repeated measures that spanned the 9-wk experimen-
tal period (F19,247 = 13.327, p = 0.001). Similarly,
across the six startle response replicates, we found sig-
nificant repeatability in female activity levels
(F19,95 = 2.539, p = 0.002: see Fig. 5), average female
distance from refuge (F19,95 = 2.65, p < 0.01), and
latency to leave refuge posture (F19,95 = 2.045,
p = 0.013).
As for startle + prey trials, we found significant
repeatability across four repeated trials in latency to
prey contact (F19,57 = 2.843, p = 0.001). However, in
these same trials repeatability was not found in female
activity levels (F19,57 = 0.875, p = 0.612), average
female distance from refuge (F19,57 = 0.883,
p = 0.604), or latency to leave refuge posture
(F19,95 = 0.981, p = 0.495). Additionally, no correla-
tion was found between these behaviors and female
body condition (Activity level: r = 0.027, N = 20,
p = 0.910, distance from refuge: r = 0.166, N = 20,
p = 0.485, Latency to leave refuge posture: r = 0.034,
N = 20, p = 0.888). Finally, in the startle + mate con-
text we found significant repeatability in female activ-
ity levels (F19,57 = 3.314, p < 0.001), average female
distance from refuge (F19,57 = 3.533, p = 0.001),
latency to leave refuge posture (F19,57 = 2.207,
p = 0.011), and female receptivity (as determined by
percentage of male activity) for both desert
(F9,27 = 3.191, p = 0.009) and urban (F9,27 = 3.133,
p = 0.01) groups (urban and desert spiders were ana-
lyzed separately due to population differences in the
percentage of male courtship).
Regarding relative repeatability across contexts, our
results showed that for female activity levels, both
startle response and startle + mate contexts were sig-
nificantly more repeatable than the startle + prey
context (antipredator vs. foraging: z = 2.47,
p = 0.014; mating vs. foraging: z = 2.93, p = 0.003).
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R. N. Halpin & J. C. Johnson Relative Plasticity in Black Widow Spiders

Table 3: Behavioral consistency (repeatability) within contexts

Startle Startle + prey Startle + mate

Measure F p r F p r F p r

Percentage of female activity 2.539 0.002 0.606 0.875 0.612 0.142 3.314 0.001 0.698
Average female distance from refuge 2.647 0.001 0.614 0.883 0.604 0.080 3.533 0.001 0.710
Latency to leave refuge posture 2.045 0.013 0.511 0.981 0.495 0.019 2.132 0.015 0.547
Number of harassments 13.327 0.001 0.930 N/A N/A N/A N/A N/A N/A
Latency to prey contact N/A N/A N/A 2.843 0.001 0.648 N/A N/A N/A
Percentage of male activity (desert) N/A N/A N/A N/A N/A N/A 3.191 0.009 0.687
Percentage of male activity (Urban) N/A N/A N/A N/A N/A N/A 3.133 0.010 0.683

Bold values are statistically significant.

activity in the startle + prey context (r = 0.423,

Fig. 5: Individuals showed a great deal of repeatability within contexts.
No significant difference in repeatability was found
between startle response and startle + mate contexts
(z = 0.47, p = 0.638). Similarly, for the average
female distance from refuge, we found that both star-
tle response and startle + mate contexts were signifi-
cantly more repeatable than the startle + prey context
(antipredator vs. foraging: z = 2.51, p = 0.012; mating
vs. foraging: z = 3.02, p = 0.003). No significant dif-
ference in repeatability was found between startle
response and startle + mate contexts (z = 0.5,
p = 0.617). Finally, for the latency to leave refuge
posture, both startle response and startle + mate con-
texts were significantly more repeatable than the star-
tle + prey context (startle response vs. startle + prey:
z = 1.7, p = 0.045; startle + mate vs. startle + prey:
z = 1.85, p = 0.032). No significant difference in
repeatability was found between startle response and
startle + mate contexts (z = 0.15, p = 0.88).
Behavioral Syndromes
Individuals that showed high levels of activity in the
startle + mate context also showed high levels of
N = 20, p = 0.06), although this correlation was mar-
ginally non-significant. However, we found no corre-
lation between activity levels in startle + prey and
startle response contexts or in startle + mate and star-
tle response contexts (startle response vs. star-
tle + prey activity: r = 0.076, N = 20, and p = 0.76,
startle response vs. startle + mate activity: r = 0.082,
N = 20, and p = 0.74). Similarly, individuals that
located themselves further from their refuge (becom-
ing more exposed to potential threats) in the star-
tle + mate context were also found to be located
further into their webs in the startle + prey context
(r = 0.585, N = 20, p = 0.008). Startle response was
not correlated with either startle + prey or star-
tle + mate behavior in regards to average female dis-
tance from refuge (startle response vs. startle + prey
distance from refuge: r = 0.064, N = 20, p = 0.80,
startle response vs. startle + mate distance from ref-
uge: r = 0.001, N = 20, p = 0.99). Finally, we did not
find a significant difference in variance between star-
tle response and startle + prey contexts (F19,38 =
0.555, p = 0.208) or between startle response and
startle + prey mate contexts (F19,38 = 1.28, p =
0.596), indicating that the lack of correlations found
between startle response and either startle + prey or
startle + mate behavior was not the result of a lack of
variation in any of our three contexts.
Discussion
We found evidence for behavioral plasticity in the
black widow’s behavioral repertoire. This plasticity was
manifested in the dramatic shifts in mean behavior
displayed across startle response, startle + prey and
startle + mate contexts. At the same time, our results
also demonstrate positive correlations between
behaviors in startle + prey and startle + mate con-
texts. This, coupled with the behavioral repeatability
found within functional contexts, lends support for

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Relative Plasticity in Black Widow Spiders
the hypothesis that black widow behavior is, to some
extent, characterized by consistent individual varia-
tion across contexts (behavioral syndromes). Thus,
black widow behavior reminds us that context-spe-
cific behavioral plasticity and context-general behav-
ioral syndromes are not mutually exclusive and may
often co-occur within a behavioral repertoire. Below,
we discuss the implications of these findings.
Desert and Urban Differences
Our results showed that urban males spent a signifi-
cantly higher percentage of their time courting urban
females relative to desert females. This courtship bias
may be the result of urban males using chemical cues
to detect and preferentially court either: (1) females
from their same populations (urban females) or (2)
females that, due to their environmental experiences,
have higher reproductive value (as urban spiders are
exposed to a surplus of prey items it might be
expected that they present a higher reproductive
value relative to their desert counterparts). Our previ-
ous work has shown that males can use silken chemi-
cal cues from females to assess a female’s feeding
status and propensity for sexual cannibalism (Johnson
et al. 2010). In addition, Kasumovic & Andrade
(2004) showed that male black widows in the field
have the ability to differentiate between conspecific
females from different geographical locations based
on chemical cues. Our data suggest that this ability to
detect population differences persists after males have
been reared in a common laboratory environment,
and thus genetic variation may underlie this trait.
Future studies will expand on this finding by quanti-
fying courtship behavior between all possible male–
female pairings (i.e., urban pairs, desert pairs, urban
female–desert male, and desert female–urban male).
Our results indicated no other urban–desert popula-
tion differences, despite the fact that these were field-
caught spiders that had been collected from their
respective habitats only a few weeks before testing
began. Thus, we suggest two possibilities: (1) urban
and desert populations might significantly differ in
other behaviors that we did not measure here or (2)
this species possesses the ability to mold behavior
quickly to variable environments (including urban,
desert, and laboratory environments), so while they
may behave differently in their respective habitats,
we are unable to see those differences in a laboratory
setting. The latter may provide insight as to how
L. hesperus is able to thrive in human-disturbed habi-
tats. Future studies will work to quantify L. hesperus
behavior under field conditions to determine whether
1244
R. N. Halpin & J. C. Johnson
and how these varied environments may result in
behavioral differences.
Behavioral Plasticity
Our data suggest that the western black widow exhib-
its a great deal of behavioral plasticity. Specifically,
females were 45% more active in startle + prey trials
relative to startle response trials. Similarly, females
were 52% more active in startle + mate trials relative
to startle response trials. This demonstrates a classic
adaptive modulation to be active when resources are
present (prey and mates) but inactive when potential
danger alone is present. Indeed, the ability of some
organisms to modulate their behavior such that they
respond adaptively in distinct contexts (behavioral
plasticity) may be a key factor in distinguishing gener-
ally between species that thrive after HIREC and those
that become locally extinct.
While a substantial number of studies on urban
birds have begun to ground the literature in urban
behavioral ecology, more studies across a range of
urban taxa are needed to develop a comprehensive
understanding of the behavioral mechanisms (e.g.,
behavioral plasticity/rigidity) underlying the success
of urban exploiters. For instance, the data presented
here suggest that, in both urban and desert popula-
tions, the western black widow displays plasticity in
startle behavior while startle + prey and star-
tle + mate behaviors are tightly coupled. In a desert
habitat, this plasticity might be expected as predators
are abundant, resources are relatively limited, and it is
crucial to behave differently toward predators vs.
either prey or potential mates. This phenotype, how-
ever, does not provide the same benefits in an urban
habitat where spiders are less likely to encounter a
predator, yet it continues to persist. This might suggest
that selection pressures in urban habitats are not
intense enough to alter the phenotype that was bene-
ficial for ancestral desert spiders. Thus, an organism’s
success in an urban habitat may be less about the abil-
ity to shift behavior (i.e., relative plasticity) and more
determined by the relative mismatch between an
organism’s historical and current environment/phe-
notype.
Behavioral Syndromes
Our results suggest that L. hesperus displays a behav-
ioral syndrome between startle + prey and star-
tle + mate behaviors while startle response behavior
remains decoupled. The coupling of startle + prey and
startle + mate behaviors presented here is similar to
Ethology 120 (2014) 1237–1247 © 2014 Blackwell Verlag GmbH
R. N. Halpin & J. C. Johnson
behavioral syndromes found in many other spider
species. For example, a common tangle-web spider,
A. studiosus, shows strong behavioral correlations
between social behavior and foraging such that indi-
viduals that are less aggressive toward conspecifics
also show reduced aggression toward prey (Pruitt
et al. 2008). Similarly, the fishing spider, Dolomedes
triton, shows a boldness syndrome between foraging
and courtship contexts as individuals that are willing
to rapidly approach prey under predation risk are also
willing to rapidly approach a potential mate under the
same perceived risk (Johnson & Sih 2007). However,
the decoupling of black widow antipredator/startle
response behavior from startle + prey and star-
tle + mate behaviors is interesting as this is not com-
monly found among spiders that display behavioral
syndromes. For instance, the behavioral syndromes
displayed by both A. studiosus and D. triton, mentioned
above, also include antipredator behavior such that
low levels of aggression toward predators is strongly
correlated with low levels of aggression toward con-
specifics and prey in A. studiosus and high levels of
boldness in the presence of predators is correlated
with high boldness in the presence of prey and mates
in D. triton.
Despite the advantages of behavioral plasticity,
behavioral syndromes are often still prevalent in pop-
ulations following HIREC (Bokony et al., 2012).
Indeed, Pintor et al. (2008) suggest that invasive cray-
fish often outcompete native species and dominate
disturbed habitats, in part, because they exhibit a cor-
related suite of highly aggressive traits (e.g., foraging
voracity, heterospecific interference). Thus, while
plasticity may often play a key role in urban exploita-
tion, adaptive behavioral syndromes may be equally
important. Future work should target potentially
non-adaptive contexts where urban exploiters would
be selected to modulate behavior. For example, the
present data illustrate a potentially non-adaptive
behavioral correlation between spider activity in star-
tle + prey and startle + mate contexts as high activity
while foraging is likely to have few costs and great
benefits while high activity while mating has been
shown to deter potential mates. The decoupling of
black widow antipredator/startle behavior from this
syndrome is consistent with the idea that selection
may favor plasticity to the extent that it allows indi-
viduals to exhibit adaptive levels of shyness to risk.
In contrast, the house sparrow, Passer domesticus,
displays a positive behavioral correlation between
risk-taking and activity levels (Bokony et al., 2012).
This correlation may be adaptive for urban birds if
high levels of activity are required to maximize forag-
Ethology 120 (2014) 1237–1247 © 2014 Blackwell Verlag GmbH
Relative Plasticity in Black Widow Spiders
ing success and the potential mortality costs of the
correlated risk-taking trait are not experienced in
urban environments that often lack key predators
(Anderies et al. 2007). We predicted that urban spi-
ders would be bolder relative to those from desert
populations because urban habitats may lack preda-
tion risk. Instead, we found no behavioral differences
between urban and desert female black widows, and a
plastic reduction of activity in the startle response
context—suggesting that, regardless of habitat, anti-
predator/startle behavior is under strong selection
(sensu life-dinner principle; Dawkins & Krebs 1979).
In conclusion, while both plasticity and behav-
ioral syndromes present distinct benefits to urban
exploiters, the presence of one within an organ-
ism’s behavioral repertoire does not exclude the
presence of the other. This is illustrated by our
results indicating that the western black widow’s
behavioral phenotype is characterized by behavioral
correlations between startle + prey and star-
tle + mate behaviors, but plasticity in startle
responses. Thus, it may be uncommon for any spe-
cies to exhibit a phenotype dominated by behav-
ioral syndromes to the exclusion of plasticity (and
vice versa), and this complexity may explain why
some species thrive following HIREC.
Acknowledgements
We would like to thank our anonymous referees for
comments on this manuscript. We also thank Theresa
Gburek who helped in collecting data. Finally, Finan-
cial assistance was provided by the National Science
Foundation (NSF-CAP3: BCS-1026865), Arizona
State University’s New College of Interdisciplinary
Arts and Sciences, the New College Undergraduate
Inquiry and Research Experiences Program, and Bar-
rett, The Honors College.
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