Animal Behaviour 186 (2022) 207e217

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Animal Behaviour
journal homepage: www.elsevier.com/locate/anbehav

Dispersal and life history of brown widow spiders in dated invasive

populations on two continents
Monica A. Mowery a, *, 1 , Yael Lubin b , Ally Harari c, Andrew C. Mason a ,
Maydianne C. B. Andrade a
a
Department of Biological Sciences, University of Toronto Scarborough, Scarborough, Ontario, Canada
b
Mitrani Department of Desert Ecology, Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev, Midreshet Ben-Gurion, Israel
c
Entomology, Volcani Center, Bet Dagan, Israel

a r t i c l e i n f o
Theory and empirical work suggest that behaviours such as dispersal and exploration are predictors of
Article history: invasive success, and that behaviours may shift predictably after invasive populations have established
Received 9 March 2021 and spread. However, there are limited data on temporal patterns in the distribution of behavioural traits
Initial acceptance 12 April 2021 linked to the timeline of establishment of invasive species. We examined dispersal and exploration, along
Final acceptance 5 January 2022 with life history traits that may be linked to behaviour, across multiple invasive populations of the brown
Available online 14 March 2022 widow spider, Latrodectus geometricus. This global invader has established populations across the United
MS. number: 21-00164R States and Israel. Using this temporal and spatial variation, we tested predictions about changes in suites
of traits over establishment time. We compared trait distributions of four U.S. populations of
Keywords: L. geometricus to patterns in four populations in Israel. We predicted that selective filters during the
behaviour
invasion process would result in more dispersive, more exploratory spiders that are larger and more
dispersal
fecund in recently established populations, but, if trade-offs occur, dispersal would be favoured at the
invasion biology
Latrodectus
expense of fecundity and size in recent populations. We found more frequent and faster dispersal in
life history more recently established populations in Israel, but not the United States. Spiders in more recently
spider established populations in Israel were larger than those in older populations, but there were no
consistent patterns across U.S. populations. However, there was evidence in both the U.S. and Israel for
differing trade-offs among fecundity, dispersal and size. In more recently established populations, spi-
ders were more variable in resource allocation to eggsacs. The results suggest that in some populations,
trade-offs underlying dispersal, fecundity and body size are shaped by the time interval, and thus the
number of generations, since establishment, with implications for the role of evolutionary processes in
invasion success.
© 2022 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

The establishment of invasive species is accelerating as a result
of climate change and human-mediated movement, with global
economic and environmental consequences (Hulme, 2009). Species
introductions do not always result in successful invasion, and our
understanding of the factors that determine successful establish-
ment is still developing (Aldorfova
et al., 2020; Kan
 uch et al., 2020;
Rehage et al., 2020; Roques et al., 2016). Establishment requires that
species are able to reproduce successfully in novel environments
(Sakai et al., 2001) in the short term, and successful invasion re-
quires adaptation to these novel conditions in the long term
* Corresponding author.
E-mail address: monicaan@post.bgu.ac.il (M. A. Mowery).
1
Present address: Mitrani Department of Desert Ecology, Ben-Gurion University
of the Negev, Blaustein Institutes for Desert Research, Midreshet Ben-Gurion, Israel.
(Blackburn et al., 2011). Initially, invasive populations may be
composed of individuals with suites of traits that allow exploitation
of a new environment, and this may include combinations of
behavioural, physiological and morphological traits not commonly
seen in the native population (Beckmann & Shine, 2011; Turner
et al., 2015). As invasive populations become better established,
traits that allowed the initial invasion may decline in frequency,
relative to adaptations to the new ecological context (Duckworth &
Badyaev, 2007).
Traits increasing invasive establishment and subsequent spread
may include larger size and high fecundity (Abhilasha & Joshi,
2009; Moravcov
a et al., 2015; Rehage & Sih, 2004), as well as
suites of behavioural traits such as increased dispersal propensity,
boldness, aggression or exploratory tendencies that can increase
movement into a new environment (Cote et al., 2010; Fogarty et al.,

https://doi.org/10.1016/j.anbehav.2022.02.006
0003-3472/© 2022 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
208 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217
2011). Correlations among behavioural traits may also have impli-
cations for ecological interactions (Canestrelli et al., 2016; Sih et al.,
2004) that could affect invasion. For example, bolder, more
dispersive and aggressive individuals consume more food and have
increased interactions with other species, and thus these in-
dividuals in relatively new invasive populations may be better
competitors (Iacarella et al., 2015; Pintor et al., 2008). Dispersal
propensity has a central importance to invasion as individuals
move past the point of introduction to establish new populations.
Dispersal tendencies may be high at the invasion front because of
spatial sorting during spread, as the most dispersive individuals
colonize new habitats and reproduce with others on the front
(Ochocki & Miller, 2017; Weiss-Lehman et al., 2017). However,
recent models of the temporal dynamics of colonization predict
that dispersal should decrease over time as an invasive population
becomes established because of a trade-off between dispersal
ability and fecundity (Perkins et al., 2013).
Although dispersal is critical to invasive success, investment in
increased dispersal ability can come at a physiological cost,
particularly in relatively new invasive populations (Ricklefs &
Wikelski, 2002; Ardia, 2005; Bonte et al. 2012). Resource con-
straints in new habitats may result in shifts in energetic allocation
to life history and other traits in populations in which high
dispersal ability is favoured (Chuang & Peterson, 2016; Hughes
et al., 2003). Relative to native populations, novel trade-offs may
arise in invasive populations, with costs manifested in concurrent
detrimental changes in other traits (e.g. Roff & Fairbairn, 2007). For
example, several studies on trade-offs at invasion fronts have found
that increased growth or faster maturation in these populations
comes at the cost of survival (Amundsen et al., 2012; Gunnarsson
et al., 2012; Hanski et al., 2006). However, trade-offs are not al-
ways predictable. For example, meta-analyses of insect species
show that investment in dispersal may co-occur with higher
fecundity (Guerra 2011; Tigreros & Davidowitz, 2019). Trade-offs
may depend on selection on life history and the costs and bene-
fits of dispersal. For instance, early juvenile dispersal could be
adaptive because of the costs of remaining at high density if siblings
cannibalize each other (Johnson et al., 2015). The varied ecological
and taxonomic patterns might be partially explained by investi-
gating patterns of dispersal and life history trade-offs across mul-
tiple independent invasion fronts of the same species, for example,
as in laboratory studies of experimental range expansion
(Fronhofer et al., 2017; Lustenhouwer et al., 2019; Van Petegem
et al., 2018).
An opportunity to infer general patterns underlying suites of
traits associated with invasion success is provided by studying
globally distributed invasive species, particularly when the timing
of invasion is known, since multiple invasive populations may be
more easily compared if they share a common genetic back-
ground. Globally distributed invasive species must survive in a
broad range of habitats, and this ecological diversity combined
with a common genetic background may enable one to distinguish
general patterns from local effects. Despite environmental het-
erogeneity, selective filtering through the stages of the invasion
process is predicted to result in similar temporal patterns in key
phenotypic traits worldwide, even while other processes occur
(e.g. kin competition, adaptation to local climate; Van Petegem
et al., 2018; Wolz et al., 2020). However, few studies have inves-
tigated patterns of trait change across a broad geographical range
to determine whether this is the case. Exceptions include a
comparative study of five native - invasive species pairs of marine
invertebrates tested in field conditions from Trinidad to New
Zealand. Across these pairs, invasive species were found to be
more stress tolerant (Lenz et al., 2011). Similarly, a study of Eu-
ropean green crabs, Carcinus maenas, showed that five invasive
North American populations were more thermally tolerant than
two populations in its native European habitat (Tepolt & Somero,
2014). A recent comparison of multiple invasive (European) and
native (Mediterranean) populations of orb-weaving spiders
showed that populations at the invasion front invested more in
reproduction but had lower hatching success, although there was
no overall difference in dispersal rates (Wolz et al., 2020). How-
ever, patterns of change in traits over invasion establishment time
have not been investigated.
In this study, we measured differences between invasive pop-
ulations in the brown widow spider, Latrodectus geometricus, a
species with a global distribution and multiple invasive pop-
ulations. The brown widow spider is a cobweb weaver (family
Theridiidae) with neurotoxic venom, purportedly native to South
Africa, and successfully established worldwide with known dates
of introduction for multiple invasive populations (Garb et al.,
2004; Taucare-Ríos et al., 2016). We focused on two indepen-
dent invasions (Israel, United States), and made direct compari-
sons between populations that were recently established and
expanding in a new environment and those with a longer history
of establishment. This species provides an excellent opportunity to
study the invasion process because of their high dispersal capacity,
ability to survive under a wide range of conditions, high repro-
ductive output and relatively short generation time (Levy &
Amitai, 1983).
We assessed a suite of traits across multiple invasive pop-
ulations, including dispersal and exploratory behaviour, body size
and life history traits that may be important to invasion (fecundity,
egg mass investment). We tested the hypothesis that behavioural
trait distributions and trade-offs among body size, fecundity and
behaviour would change over population establishment time as a
function of the stage of invasion (Blackburn et al., 2011). Specif-
ically, we predicted that individuals from newly established pop-
ulations would be more dispersive compared to individuals from
longer established populations. Finally, we assessed trade-offs
among investment in dispersal, fecundity and body size by exam-
ining correlations within individuals and family lines in Israel and
the United States.
The general pattern of trade-offs with dispersal is that invest-
ment in dispersal would result in less investment in fecundity (e.g.
Duthie et al., 2015). However, in spiders, dispersal of recently
emerged young is by ballooning; this is a passive process and does
not require investment in costly phenotypic traits often needed for
long-distance locomotion. Consequently, we would not expect a
trade-off between dispersal and adult body size or fecundity.
Dispersal of invasive widow spiders may be largely human medi-
ated (e.g. transport by cargo ships, Nentwig 2015). Adult females
with large body size will be able to survive without food over long
periods of transport and post-transport establishment (Forster and
Kavale, 1989). Large female body size is also positively related to
fecundity: larger females produce more eggs rather than larger
eggs, while egg size appears to be species specific (Anderson, 1990).
Given the relationship between size and fecundity, an alternative
prediction would be that dispersal of adult females will trade off
with fecundity.
If similar correlations are pervasive across both invasions, it
suggests that allocation to one trait comes at the cost of the other
(e.g. female dispersal and fecundity), and these relationships may
impose constraints on changes during establishment. In contrast, if
the trade-offs vary between populations or with time since estab-
lishment, then correlations may arise from variation in allocation
tactics, or adaptive links between traits. This work is novel because
we examined changes in invasive behaviours in concert with life
history and behavioural trade-offs across time in dated invasions on
two continents.
 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217
METHODS
Study Species
Brown widow spiders were first reported in North America in
southern Florida in 1935 (Pearson, 1936). By the 1960s,
L. geometricus was abundant in coastal cities in southern Florida,
but was rarely found in surveys in the northern part of the state
(Levi, 1959; McCrone & Stone, 1965). Since that time, these spiders
have spread to Texas, where they were first detected in 1997,
although a collection location was not specified (Garb et al., 2004).
Brown widows were first detected in South Carolina in 2001
(Howard, 2017). They have expanded their range across the
southern United States, and are found sporadically in New Mexico
and southern Arizona, according to records from citizen scientists
(research-grade verified records, iNaturalist, https://www.
inaturalist.org). In the west, L. geometricus was first found in Tor-
rance, California (Los Angeles County) and Orange County in 2003
and 2004, confirmed by collections in the Los Angeles County
Natural History Museum and specimens submitted to Orange
County Vector Control (Vincent et al., 2009). In 2006, Fullerton
College (Fullerton, CA) arachnology students collected
L. geometricus specimens for the first time, despite yearly collec-
tions starting in 1996 (Vincent et al., 2009). By 2008, L. geometricus
was found in Los Angeles County, Orange County, San Diego County
and Riverside County (Vincent et al., 2009). Human-mediated
transport and multiple invasions are likely have affected spread
because of high connectivity, despite the thousands of kilometres
between populations (Banks et al., 2015). In particular, because of
the short time frame of the spread of L. geometricus to California, it
is likely that spiders were either moved through human-mediated
transport or that the California population was from a novel
introduction.
A more recently established invasive population is in Israel,
where L. geometricus was first detected in Tel Aviv in 1980. The
species was not detected in a taxonomic study of the genus Latro-
dectus in 1966 (Levi, 1966; Levy & Amitai, 1983). Spiders have since
spread south into the Negev Desert, where they have been estab-
lished since the early 2000s (Y. Lubin, personal observation; see
Fig. 1b for a map of invasive populations in Israel). Latrodectus
geometricus was first observed in Beer Sheva in 2000 and spread to
Yeruham and then Sede Boqer several years later (Y. Lubin, personal
observation). The populations across Israel are separated by
100e200 km and rural populations in the Negev Desert are less
likely to be highly interconnected, which reduces the likelihood of
introductions from multiple sources. We use this variation in in-
vasion history to make a systematic comparison of traits along a
gradient from established invasive populations to the invasion
front.
Latrodectus geometricus spiders typically build their webs in
habitats associated with humans, including fences, parks, children's
playgrounds and bridges (Vincent et al., 2009). These urban habi-
tats are more similar in climate and physical conditions (e.g. urban
areas across the U.S. (Hall et al., 2016), and often have more prey,
than surrounding natural environments (Lowe et al., 2016). Even
across environments with cold winters or low precipitation, the
extreme conditions in these areas that could be unsuitable for
L. geometricus survival are mitigated by the urban heat island effect
(Johnson et al., 2019) and high humidity due to irrigation (Cook &
Faeth, 2006).
Brown widow females are extraordinarily fecund. Mated fe-
males store sperm and can produce eggsacs containing between 50
and 200 fertilized eggs (Danielsen et al., 2014) every few days if
well fed and for up to 2 years after a single mating (Waner et al.,
2018). Spiderlings hatch inside the eggsac and moult once before
209
emerging as second instars (Kaston, 1970), then stay on their natal
web for a week, during which time they often cannibalize siblings.
Spiderlings then disperse by walking, rappelling (short range) or
ballooning (long range) to new habitats (Johnson et al., 2015) where
they must build their own capture webs to catch prey. Offspring
dispersal propensity is likely to be influenced by the maternal
environment (Mestre & Bonte, 2012).
Populations are typically multivoltine with spiders of all
developmental stages present through much of the season
(Andrade & MacLeod, 2015 and references therein). In other
Latrodectus species, mated adult females, later-stage juveniles and
penultimate instar males can overwinter. Females become sexually
mature after 4 months (Heeres & Clark, 1997), depending on food
availability and season. An invasive population of brown widows
that has been established since 1935, such as in Florida, will have
undergone between 200 and 250 generations by 2019, whereas
more recently established populations, such as those from the early
2000s (e.g. the Sede Boqer and Los Angeles populations) will have
undergone 20e60 generations.
Study Populations
Mated female spiders and eggsacs were collected from sites
across the dated invasion in the United States (Fig. 1a; Gainesville,
Florida: 29.65
N, 82.34
W; Edisto Beach Park, South Carolina:
32.51
N, 80.30
W; Austin, Texas: 30.28
N, 97.75
W; Riverside,
California: 33.98
N, 117.33
W) and Israel (Fig. 1b; Tel Aviv: 32.10
N,
34.80
E; Beer Sheva: 31.26
N, 34.80
E; Yeruham: 30.98
N, 34.93
E;
Sede Boqer: 30.85
N, 34.79
E). Spiders from the Israel populations
were reared on the Sede Boqer campus of Ben-Gurion University of
the Negev. Spiders collected in the United States were shipped to
the University of Toronto Scarborough where they were reared in
the Invertebrate Rearing Facility. All spiders from the U.S. used in
these experiments were reared in the laboratory under 12:12 h
light:dark and at 25
C.
Field-collected adult female spiders in Israel were fed one
20 mm long grasshopper nymph, Schistocerca gregaria, per week
and reared at 26 ± 1
C, 65e70% relative humidity and 14:10 h
light:dark for 1 month before we assessed body size and offspring
traits (from eggsacs produced in the laboratory after this period).
Adult female spiders from the United States populations were fed a
20 mm long cricket (Gryllodes sigillatus or Acheta domesticus) per
week and kept at 12:12 h light:dark and 25
C for 1 month before
we similarly assessed size and offspring traits. For reasons of
availability, spiders from the U.S. and Israel received different prey
species with possibly different nutrient content, which could have
affected some behaviours (Koemel et al., 2019). Field-collected fe-
males were checked every other day for newly deposited eggsacs,
which were collected and weighed, and placed in separate plastic
vials (2 cm in diameter x 6 cm). Sacs were checked daily for spi-
derling emergence and the date was recorded. Spiderlings were
counted and photographed, with size and behaviour (exploration
and dispersal) measured 1 week after emergence, prior to which
spiderlings were not fed or given water (typical for spiderlings prior
to dispersal).
Behaviour
We measured three behavioural traits relevant to invasive
spread. (1) Dispersal propensity was the proportion of spiderlings
within a family line that dispersed by rappelling or ballooning in
our trials (see below). (2) Dispersal time, measured only for spi-
derlings that dispersed by rappelling or ballooning, was the interval
between the trial start to the spiderling's arrival at the end wall of
the dispersal arena. (3) Exploratory behaviour was the latency
210 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217
50°N (a)
45°N
Latitude
40°N
35°N Riverside, CA
Austin, TX
30°N
1000 km
25°N
130°W 120°W 110°W 100°W
Figure 1. Map of the study populations (marked in red) of Latrodectus geometricus in (a) the United States and (b) Israel.
(time) for a spiderling to commence movement and lay down silk
after being placed in a novel environment, a behaviour that might
relate to the speed of postdispersal establishment of a web.
Dispersal was measured in spiderlings that were offspring of
field-mated adult females, one eggsac per female, with five spi-
derlings tested from each family line (number of family lines: US:
Florida N ¼ 4, Texas N ¼ 6, South Carolina N ¼ 12, California N ¼ 22;
Israel: Tel Aviv N ¼ 24, Beer Sheva N ¼ 5, Yeruham N ¼ 14, Sede
Boqer N ¼ 10). Measurements were made 7 days after emergence
from the eggsac, which is a typical interval before dispersal
(Johnson et al., 2015). Spiderlings were held communally in sibling
groups prior to that time.
Our dispersal assay was similar to that reported in Johnson et al.
(2015). We placed individual spiderlings on a wooden dowel inside
a rectangular Plexiglas arena (0.50 x 0.34 m and 0.35 m high) held
at 23
C. One end wall was replaced by a mesh screen and the other
end had a mesh-covered circular hole through which a fan blew air
at 1 m/s. After a spiderling settled on the dowel, we turned the fan
on to start the trial, which ended after 10 min. We recorded
whether the spider remained on the dowel, rappelled away from
the dowel (left the dowel while anchored by a dragline) or bal-
looned away from the dowel (tilted the posterior abdomen up-
wards, released silk into the air such that it was caught by the wind
so the spider ‘flew’ away from the dowel, Bonte et al., 2003).
Dispersal propensity was measured as a binary variable (yes or no)
and dispersal time as the minutes elapsed from the trial start to
when the spider reached the mesh wall, 0.29 m from the dowel,
after ballooning or rappelling.
Exploratory behaviour was measured as the latency to initiate
movement and silk laying in a new container for each of five spi-
derlings per eggsac, one sac per female (family lines: Tel Aviv
N ¼ 17, Beer Sheva N ¼ 5, Yeruham N ¼ 15, Sede Boqer N ¼ 8).
Exploratory behaviour was measured for two U.S. populations,
Texas (family lines, N ¼ 7) and South Carolina (N ¼ 8). After
measuring dispersal propensity, each spiderling was placed in an
empty, unfamiliar plastic cage (5 x 5 cm and 7 cm high, clear plastic
container, Amac Plastics, Ltd., Petaluma, CA, U.S.A.) and the time
until it started to move and release silk was measured. When spi-
ders move, they release a silk dragline behind them (Foelix, 2011),
so initiation of movement and silk release coincided.
Life History
Fecundity was quantified as the total number of eggs per sac
(emerged spiderlings þ unhatched eggs left in the sac) and
32.5°N (b)
32.0°N Tel Aviv
31.5°N
Beer Sheva
Edisto Island, SC
31.0°N Yeruham
Sede Boqer
Gainesville, FL
30.5°N 50 km
90°W 80°W 70°W 34.0°E 34.5°E 35.0°E 35.5°E 36.0°E
Longitude
reproductive investment as mass of the sac. For the Israel pop-
ulations, fecundity was assessed in the first eggsac produced by
each female in the laboratory (Tel Aviv N ¼ 45, Beer Sheva N ¼ 18,
Yeruham N ¼ 39, Sede Boqer N ¼ 20), and mass was measured
shortly after sac production using a balance (Mettler ME204). For
the United States populations, fecundity was assessed from eggsacs
collected in the field prior to hatching and brought into the labo-
ratory (Florida N ¼ 11, Texas N ¼ 7, South Carolina N ¼ 13, California
N ¼ 25).
One eggsac per female was randomly selected per nest to avoid
pseudoreplication of using multiple fecundity estimates per indi-
vidual. Since eggsacs were collected from different webs at each
site, these represent fecundity measures for different females. For
field-collected eggsacs, eggsac order was unknown. The variation in
number of eggs per sac correlates with female body size in many
spiders (Anderson, 1990), and in a congener, the Australian redback
spider, Latrodectus hasselti, in which lifetime reproduction was
assessed in laboratory-reared females, eggsac yields (number of
spiderlings) began to decline below average only after 15 eggsacs
were produced and females were beginning to senesce (Andrade &
Banta 2002). In nature, L. hasselti females produce an average of five
eggsacs in their lifetime (Andrade & Banta 2002). Taken together,
this suggests that choosing one of the female's eggsacs can give a
reasonable picture of reproductive potential. Eggsac mass was not
recorded since sacs were produced in the field.
Body Size
We measured the body size of adult females as well as the size of
juveniles at the second instar from digital photographs (Nikon DXM
1200 digital camera mounted on a Zeiss Stemi 2000-C dissecting
microscope), using ImageJ (Abramoff et al., 2003). We measured
live adult field-collected females (patella - tibia length, Jakob et al.,
1996) from Tel Aviv (N ¼ 30), Beer Sheva (N ¼ 21), Yeruham
(N ¼ 37), Sede Boqer (N ¼ 20), Florida (N ¼ 16), Texas (N ¼ 11),
South Carolina (N ¼ 20) and California (N ¼ 20). Females were
photographed while restrained in a clear plastic bag, with their legs
held flat on cotton batting; all spiders survived this procedure with
no apparent ill effects. The body size of juveniles (width of the
cephalothorax, Jakob et al., 1996) was measured at the second
instar, immediately after their emergence from the eggsac, based
on digital photographs of 10 spiderlings from each eggsac (U.S.
family lines: Florida: N ¼ 4, South Carolina N ¼ 9, Texas N ¼ 7,
California N ¼ 6; Israel family lines: Tel Aviv, N ¼ 13 and Sede Boqer,
N ¼ 2).
 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217
Statistical Analysis
We analysed the data separately for the four Israel populations
and for the four populations from the United States. We did not
analyse the eight populations together along a quantitative time-
scale of invasion establishment because the differences in estab-
lishment time were orders of magnitude different, and the
establishment times for the Beer Sheva, Yeruham and Sede Boqer
populations were approximate.
To compare adult body size and fecundity along the dated
invasions, we used a one-way ANOVA by country with population
as the predictor variable, followed by post hoc Tukey tests. Dif-
ferences in spiderling traits (second-instar spiderling size,
exploratory behaviour and dispersal time) were assessed using
general linear mixed-effects models (GLMMs) for each target
variable as a response, family line (random factor) and population
(fixed factor) as predictors, Gaussian link functions for body size,
fecundity, eggsac mass and dispersal time. For exploratory
behaviour, a Poisson link function was used. For binary dispersal
propensities, we used a general linear mixed model (GLM) with
predictors including family line (random factor), population (fixed
factor) and dispersal as the response variable with a binomial
distribution.
To test for links between invasion history, exploratory behav-
iour and dispersal tendency, we used a GLMM with dispersal
propensity as the binary response variable, latency to explore and
invasion establishment time as predictors and family line as a
random factor. To assess the relationship between adult female
body size and fecundity, we used a GLM with fecundity as the
response variable, and body size and invasion establishment time
as predictors.
We examined whether trade-offs differed between longer
established and more recently established populations. Trade-offs
would be suggested by significant positive or negative relation-
ships between variables (Roff & Fairbairn, 2007). To test for dif-
ferences in correlations between maternal patella - tibia length
and fecundity, we created a GLMM with fecundity as the response
variable, and invasion establishment and maternal patella - tibia
length as the predictors. To test for effects of invasion establish-
ment time on the relationship between eggsac mass and number
of eggs per eggsac, we created a GLMM with fecundity as
the response variable, and invasion establishment time and egg-
sac mass as predictors, including an interaction between
predictors.
All statistics were performed using R version 3.3.2 (R Core Team,
2020). GLMMs used the package ‘lme4’ (Bates et al., 2015) and plots
were created using ‘ggplot2’ (Wickham et al., 2019).
Ethical Note
We collected L. geometricus adult female spiders and eggsacs
for this study in sites across established invasive populations in
the United States and Israel. Spiders were collected from road-
sides, public areas and around buildings with permission of
landowners. In these sites, L. geometricus was well established and
collecting was unlikely to affect population size. The spiders were
held in the laboratory in accordance with the ASAB/ABS Guidelines
for the treatment of animals in behavioural research and teaching.
Behavioural trials and size measurements were not harmful to the
assayed individuals. Spiders were reared individually after the
third instar to reduce cannibalism and, after behavioural mea-
sures, were kept to the end of their natural life spans in the
laboratory.
211
RESULTS
Behaviour
Recently established Israel populations in Yeruham and Sede
Boqer had higher dispersal propensity than the longer established Tel
Aviv and intermediate Beer Sheva populations (Table 1, Fig. 2a). Spi-
derlings from recently established populations (Beer Sheva, Yeruham
and Sede Boqer) were also faster to disperse than spiderlings from the
long-established Tel Aviv population (Table 1, Fig. 2b). In the United
States, there was significant variation in dispersal propensity, driven
primarily by low dispersal in the South Carolina population (inter-
mediate establishment time). The recently established California
population was more likely to disperse than the longer established
South Carolina population (Table 1, Fig. 2c). However, contrary to our
predictions, spiderlings from the long-established Florida population
were also more likely to disperse than the South Carolina population.
In terms of dispersal time, there was considerable variation, and no
significant differences across populations, although spiderlings from
California tended to have shorter times to disperse than those from
South Carolina (Table 1, Fig. 2d).
Exploratory behaviour in a new environment was slower in the
intermediate Beer Sheva population than the Tel Aviv and Yeruham
populations (Table 1). In the United States populations, spiderlings
from Texas were significantly faster to explore than the South
Carolina population (Table 1). Exploration and dispersal were
positively correlated, although behavioural correlations did not
differ with invasion establishment time. Across populations from
the U.S. and Israel, spiders that were faster to explore were more
likely to disperse across both the longer and recently established
populations, with no interaction between exploration and invasion
establishment time (GLMM: invasion: z ¼ 3.129, P ¼ 0.002; latency
to explore: z ¼ -3.870, P < 0.001).
Life History
To assess differential allocation to reproduction as a function of
invasion progression, we tested for differences in fecundity within
Israel and U.S. populations. Populations in Israel had consistently
low fecundity (Table 1, Fig. 3). In contrast, fecundity varied signif-
icantly across populations in the United States (Table 1, Fig. 3), with
the median number of eggs ranging from 50 eggs per sac in Cali-
fornia to almost 150 eggs per sac in Florida. The most recently
established California population was less fecund than longer
established Florida, Texas and South Carolina populations.
Across both the U.S. and Israel, larger mothers produced eggsacs
containing more offspring and country of origin also affected the
number of eggs in an eggsac, but the fecundity of spiders from the
recently established populations was not significantly different for
their body size than that of similarly sized spiders from longer
established sites (country: t ¼ 4.830, P < 0.001; maternal size:
t ¼ 2.905, P ¼ 0.004; invasion establishment: t ¼ 1.206, P ¼ 0.231).
However, for any given increase in maternal allocation (eggsac
mass), the number of eggs in the sac increased at a faster rate in the
Yeruham than in the Tel Aviv population in Israel (Fig. 4; GLM:
interaction between eggsac mass and population: t ¼ 3.306,
P ¼ 0.002). We had insufficient data to include a similar analysis
from the United States.
Body Size
Adult body size varied significantly across Israel invasive pop-
ulations and across populations from the United States. In the
212 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217

Table 1
Results for models of dispersal behaviour, exploration, fecundity and size in L. geometricus across populations in the U.S. and Israel

Trait Model: U.S. df, P Model: Israel df, P

Dispersal propensity GLMM: 3, 261 GLMM: 3, 222

F ¼ 5.070 F ¼ 2.753
Dispersal propensity: CA-SC P ¼ 0.043 Tel Aviv-Yeruham z ¼ 3.424 P < 0.001
pairwise comparison z ¼ -2.020 Tel Aviv-Sede Boqer z ¼ 2.583 P ¼ 0.009
Dispersal time GLMM: 3, 97 GLMM: 3, 170
F ¼ 1.727 P < 0.001 F ¼ 6.129 P ¼ 0.08
Dispersal time: NS Tel Aviv-Beer Sheva t ¼ 3.297,
pairwise Tel Aviv-Yeruham P < 0.001
comparisons Tel Aviv-Sede Boqer t ¼ 3.361,
P < 0.001
t ¼ 2.828,
P ¼ 0.005
Exploratory behaviour GLMM: 2, 39 GLMM: 3, 215
z ¼ 3.848 P < 0.001 z ¼ 2.318 P ¼ 0.021
Exploratory behaviour: TX-SC t ¼ -1.696 Beer Sheva-Tel Aviv t ¼ -1.860
pairwise comparison Beer Sheva-Yeruham t ¼ -1.623
Fecundity ANOVA: 3, 52 ANOVA, 3, 118
F ¼ 12.18 P < 0.001 F ¼ 1.008 P ¼ 0.392
Fecundity: pairwise CA-TX P ¼ 0.014 NS
comparison CA-SC P ¼ 0.002
CA-FL P < 0.001
Adult female size ANOVA: 3, 63 ANOVA: 3, 72
F ¼ 12.26 P < 0.001 F ¼ 7.208 P < 0.001
Adult female size: SC-FL P < 0.001 Tel Aviv-Beer Sheva t ¼ 3.115,
pairwise comparison SC-TX P < 0.001 Tel Aviv-Yeruham P ¼ 0.012
SC-CA P < 0.001 Tel Aviv-Sede Boqer t ¼ 3.064,
P ¼ 0.014
t ¼ 4.359,
P < 0.001
Juvenile size LMM: 3, 492 LMM, 3, 138
F ¼ 1.579 F ¼ 0.011
Juvenile size: pairwise NS NS
comparison

invasive populations in Israel, despite considerable variation, field-
collected adult female spiders from more recently established
populations were larger than those from Tel Aviv, the earliest-
established site (Table 1, Fig. 5a). This effect was significant for
spiders from Beer Sheva, Yeruham, and Sede Boqer, which were all
significantly larger than spiders from the longer established Tel
Aviv population. Field-collected adult females from United States
populations (Florida, South Carolina and Texas) also differed in size
(Table 1, Fig. 5b). Adult females from an intermediate established
population from South Carolina were smaller than those from
Florida, Texas and California. Offspring of field-collected females
measured as juveniles, however, did not differ in size across either
invasion (Table 1).
DISCUSSION
The worldwide distribution of invasive L. geometricus spiders,
and their well-documented invasion history, allowed us to test for
shifts in phenotypic traits by sampling across invasive populations
in Israel and the United States. We measured some of the traits
thought to be key to invasion success, including dispersal, explor-
atory behaviour, reproductive investment and body size, in several
longer established and recently established populations. Recently
established populations should have spiders that are highly
exploratory and dispersive, but this may come at the cost of
reproductive investment (Duthie et al., 2015). In longer established
populations, the dispersive and exploratory tendencies should
decrease with a concomitant increase in fecundity, which, for in-
vertebrates, may partly arise from increased body size.
We found higher dispersal rates and shorter latency to disperse
in more recently established invasive populations in Israel (Fig. 2a
and b). In the United States, however, there was no pattern in
dispersal propensity across the invasion front (Fig. 2c and d).
Dispersal propensity may be favoured at range edges in recent in-
troductions that occurred over smaller geographical ranges (i.e.
Israel). Within the United States populations, fecundity was higher
in the longer established populations, although this was not the
case in Israel (Fig. 3). Populations showed different relationships
between egg investment and fecundity (Fig. 4). Field-collected
adult females were larger in recently established populations in
Israel (Fig. 5a), although recently emerged juveniles were not
different in size.
Spiderling dispersal (ballooning and rappelling) is critical for
invasive spread following establishment. While the Israel pop-
ulations had greater dispersal tendency and speed in recently
invaded areas, as predicted, this was not the case in the U.S. pop-
ulations. Individuals in invasive populations of L. hasselti are more
likely and faster to disperse than those in a native population
(Mowery et al., 2021), suggesting that dispersal could be linked to
invasiveness in other widow spider species. Across populations in
the U.S. and Israel, we found that individuals that were more
exploratory were also more likely to disperse. These behaviours
may be linked by having generally increased activity levels, as
found in other taxa, including roe deer, Capreolus capreolus, and
fruit flies, Drosophila spp. (Debeffe et al., 2013; Tung et al., 2018).
Spiderling dispersal is mainly passive requiring little energy
expenditure (Szymkowiak et al., 2007), and is thus unlikely to be
traded against the ability to invest in future reproduction. A recent
study of invasive populations of orb-weaving spiders, Argiope
bruennichi, in Europe found no consistent patterns in dispersal
relative to recency of population establishment (Wolz et al., 2020),
and suggested that climate differences could mask dispersal and
life history shifts in populations over that invasive range. Given that
significant climatic shifts accompany the A. bruennichi invasion
 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217 213

(a) a a b b (b)
a b b b
1
600

0.8

400
0.6

0.4 200

0.2
Proportion dispersed

Time to disperse (s)

N = 24 N=5 N = 14 N = 10 0 N = 64 N = 16 N = 56 N = 38
Tel Aviv Beer Sheva Yeruham Sede Boqer Tel Aviv Beer Sheva Yeruham Sede Boqer

(c) a ab b a (d) NS
1 600

0.8

400
0.6

0.4
200
0.2

0
N=4 N=6 N = 12 N = 22 0 N = 13 N = 16 N = 17 N = 55
Florida Texas South California Florida Texas South California
Carolina Carolina
Population

Figure 2. (a, c) The propensity to disperse measured as the proportion of L. geometricus spiderlings per eggsac to disperse in a 10 min testing period. Sample size is the number of
eggsacs from each population and black points represent outliers. (b, d) Time to disperse. Sample size is the number of individuals and black points represent individual spiderlings.
(a, b) Invasive Israel populations: the longer established central population (Tel Aviv), intermediate population (Beer Sheva) and recently established populations at the invasion
front in the Negev (Yeruham and Sede Boqer). (c, d) invasive United States populations: the long-established population from Florida, intermediate populations from South Carolina
and Texas and a recently established population from California. The box plots show the median and 25th and 75th percentiles; the whiskers indicate the values within 1.5 times the
interquartile range. Different letters above box plots indicate significant differences between locations based on Tukey HSD post hoc tests.

(Mediterranean to Germany), a plastic response of dispersal to
temperature could confound other effects of establishment time. In
the L. geometricus invasions described here, by comparison, climatic
shifts could be a factor affecting patterns of morphology and
fecundity, especially given the wide variation in precipitation and
temperature across the invasive ranges in Israel and the United
States.
Climate gradients across the invasive range could explain pop-
ulation level differences in traits, including body size (e.g. in spi-
ders, Entling, Schmidt-Entling, Bacher, Brandl & Nentwig, 2010). In
the invasive orb-weaver Cyrtophora citricola, a combination of local
adaptation and spatial sorting may explain population differences
in exploration, boldness and voracity across a latitudinal gradient
(Chuang & Riechert, 2021). Across our four United States sites,
which varied primarily in longitude (latitudinal range 29.65
N to
33.98
N, longitudinal range 80.30
W to 117.33
W), there were no
consistent patterns in temperature or humidity. Invasive brown
widow spiders live in urban environments, which may be similar in
microhabitat conditions and are more homogeneous in tempera-
ture and humidity than nearby natural habitats (Hall et al., 2016).
Similarly, cosmopolitan, invasive pavement ants are able to
colonize a wide range of climates by thriving in urban areas
(Cordonnier et al., 2020).
In brown widow spiders, similar to other anthropophilic species,
organism-initiated dispersal combined with human-mediated
transport likely drive population spread (Gippet et al., 2019;
Zhang et al., 2019). The combination of human-mediated and nat-
ural dispersal drives expansion of other invasive species, such as
emerald ash borers, Agrilus planipennis (Muirhead et al., 2006). In
addition to leading to more rapid population spread, human
transport will decrease the likelihood of divergence due to ongoing
introduction of new individuals. Urban populations are hubs of
connectivity, and in the increasingly globalized Anthropocene era,
geographical distance is a less significant predictor of divergence
than urbanization (Capinha et al., 2015). Patterns of trade and
transport methods could explain differences in trait distributions
between independent invasion fronts of the same species (Banks
et al., 2015), and increased connectivity may explain the lack of
patterns in the U.S. populations based on establishment time
compared to the populations in Israel.
There were differences between the U.S. and Israel in the tem-
poral shifts in traits, as well as in mean trait values and trade-offs
214 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217

300 b b b a
Fecundity (no. of eggs per sac)

200

a a a a

100

0 N = 45 N = 18 N = 39 N = 20 N = 11 N=7 N = 13 N = 25

Tel Aviv Beer Sheva Yeruham Sede Boqer Florida Texas South California
Carolina
Population

Figure 3. Fecundity (number of eggs in the first eggsac produced in the laboratory), across populations of L. geometricus in Israel (long-established Tel Aviv, intermediate Beer Sheva
and recently established Yeruham and Sede Boqer populations) and the United States (long-established Florida, intermediate South Carolina and Texas and recently established
California populations). The box plots show the median and 25th and 75th percentiles; the whiskers indicate the values within 1.5 times the interquartile range. Black points
represent individual eggsacs. Different letters above box plots indicate significant differences between locations based on Tukey HSD post hoc tests.

could be significant (e.g. Burton et al., 2010; Iacarella et al., 2015). In

150 Invasion
the United States, L. geometricus individuals may need to allocate
Tel Aviv more resources towards competition and fewer resources to other
Yeruham traits such as fecundity, or they could suffer costs that impede in-
Fecundity (no. of eggs per sac)

vasion and dispersal (Burton et al., 2010). Latrodectus mactans, the

100
50
0 other findings that larger invasion front individuals have more
0 0.025 0.05
Eggsac mass (g)
Figure 4. Regression of L. geometricus eggsac mass and number of eggs per sac in the
initial site of introduction (Tel Aviv) and in a newer established population (Yeruham).
Grey area around the regression lines represents the 95% confidence interval.
between traits, which suggests that, even within a species, trait
shifts may not be consistent. These differences may result from
different local ecological conditions. For example, in theoretical and
experimental studies, the effect of differences in local competitors
native southern widow spider, is prevalent in many of the sampled
areas in the southern United States, and it has been found to occupy
habitats where brown widows were previously seen (Garb et al.,
2004; Taucare-Ríos et al., 2016). Invasive brown widows appear
to allocate resources to fast dispersal (Fig. 2), but may be less
competitive in interspecific interactions (Jones et al., 2019; Lewis,
2013). In contrast, in Israel, while three congeners are present
(Latrodectus pallidus, Latrodectus revivensis, Latrodectus trede-
cimguttatus), there is little habitat overlap, and so competition is
less likely to influence population spread (Levy & Amitai, 1983).
Selection for larger size at the invasion front is consistent with
endurance and dispersal potential, such as in invasive cane toads,
0.075
Rhinella marina (Phillips et al., 2006), as well as higher fecundity in
gobies, Neogobius melanostomus (Brandner et al., 2013) and plants
(Jelbert et al., 2015). In our study, however, these size differences in
field-caught females could have been influenced by environmental
differences during development, such as temperature or resource
availability. Larger adult body size in the invasion front populations
was not correlated with higher fecundity. However, more recently
invasive spiders increased eggsac investment (number of offspring)
with increased maternal resources (eggsac mass) compared to
spiders from older populations. The measurement of fecundity that
 M. A. Mowery et al. / Animal Behaviour 186 (2022) 207e217 215

8 (a) a b b b 8 (b) a a b a

7 7
Patella–tibia length (mm)

6 6

5 5

4 4

3 N = 31 N = 23 N = 38 N = 21 3 N = 16 N = 11 N = 20 N = 20

Tel Aviv Beer Sheva Yeruham Sede Boqer Florida Texas South California
Carolina
Population

Figure 5. Patella - tibia length of L. geometricus adult females with population location arranged in order of decreasing age of establishment (left to right) (a) in Israel, from the long-
established central population (Tel Aviv) and intermediate population (Beer Sheva) to the recently established populations at the invasion front in the Negev (Yeruham and Sede
Boqer), and (b) in the United States, a long-established population in Florida, intermediate populations in Texas and South Carolina and a newly introduced California population.
The box plots show the median and 25th and 75th percentiles; the whiskers indicate the values within 1.5 times the interquartile range. Black points represent size of individual
female spiders. Different letters above box plots indicate significant differences between locations based on Tukey HSD post hoc tests.

we used, the number of eggs per sac, is one measure of reproduc-
tive success, since females lay multiple eggsacs over their lifetime.
Lifetime reproductive success could vary across populations
depending on both the number of eggs per sac and the rate of
eggsac production (Waner et al., 2018). For example, in L. hasselti,
spiders from an invasive population from Japan produce eggsacs
more frequently than a native population from Australia, but eggsac
contain similar numbers of eggs (Mowery et al., 2021). Fecundity of
one clutch is also variable across seasons and years depending on
shifting abiotic and biotic factors, so differences across populations
may reflect resource availability at a given point in time (Danielsen
et al., 2014). As such, assessing lifetime reproductive success, taking
the quantity, frequency and variability of reproduction into ac-
count, would provide a more complete comparison of life history
differences between invasive populations.
Overall, we found consistent patterns in larger adult body size,
high dispersal propensity and more rapid dispersal in more
recently established invasive populations in Israel, but not in U.S.
populations. There were no consistent patterns in exploratory
behaviour with invasive establishment time, but across pop-
ulations, more exploratory individuals were more likely to disperse.
Divergent trade-offs between populations indicate that dispersal is
not constrained by maternal size or fecundity. Increased plasticity
in eggsac investment may be adaptive in variable environments.
Our results support hypotheses based on the idea that selection
throughout the invasion process, and in the invaded habitats away
from the point of introduction, results in populations with different
size, life history and behaviour. Additional studies using approaches
like this one in other cosmopolitan, urban species that compare
discrete invasion fronts for a globally invasive species would in-
crease our understanding of the patterns of invasive spread and
population level changes behind invasion success.
Author Contributions
M.A.M., A.C.M., Y.L. and M.C.B.A. conceived of the study, M.A.M.
collected data with suggestions from A.H. and Y.L., M.A.M. wrote
the initial manuscript draft with critical input from A.C.M., Y.L. and
M.C.B.A. Funding was acquired by M.A.M. and M.C.B.A. All authors
approved the final version of the manuscript.
Data Availability
The data sets analysed in this study are available in the Dryad
repository at https://datadryad.org/stash/dataset/doi:10.5061.
Declaration of Interest
The authors declare they have no competing interests.
Acknowledgments
We thank C. Chapman, C. Jones, A. Fuller and N. Singh for col-
lecting spiders from the United States and C. Scott and M. Segoli for
helpful comments on the manuscript. We thank L. Sentenska for
photographs of L. geometricus from California. This study was
completed in partial fulfilment of the requirements for a PhD in the
Department of Ecology and Evolutionary Biology (University of
Toronto). M.A.M. was funded by a Mitacs Globalink fellowship and
an Animal Behavior Society George W. Barlow award. Research at
UTSC was supported by an NSERC Discovery grant (#2017-06060)
and Leaders Opportunity fund grant from the Canada Foundation
for Innovation (#203764) to M.C.B.A.
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