Behavioral Ecology and Sociobiology (2019) 73: 127

https://doi.org/10.1007/s00265-019-2741-5

ORIGINAL ARTICLE

Pay attention to the ladies: female aggressive behavior and weapon

allometry provide clues for sexual selection in freshwater anomurans
(Decapoda: Aeglidae)
Marcelo M. Dalosto 1 & Luciane Ayres-Peres 2 & Paula B. Araujo 3 & Sandro Santos 1 & Alexandre V. Palaoro 4

Received: 5 April 2019 / Revised: 1 August 2019 / Accepted: 5 August 2019 / Published online: 19 August 2019
# Springer-Verlag GmbH Germany, part of Springer Nature 2019

Abstract
Contesting scarce resources can trigger the evolution of specialized morphological structures (i.e., animal weapons). While most
research focus on male weapons, females might also bear weapons, although generally smaller and less conspicuous than male
weapons. Social selection is evoked to explain female weaponry in which females fight for nonsexual resources such as food and
shelter. Males might fight for similar resources but are expected to have proportionally larger weapons due to additional inputs from
sexual selection. We tested whether males have proportionally larger weapons than females in two species of Aegla crabs. Interestingly,
only males of one species had proportionally larger claws than females. Given that these larger weapons typically correlate to increased
aggression, we expected males to fight more intensely than females. Thus, we compared intrasexual contests of males and females of
the same species. Females fought similarly to males: latency, contest duration, and frequency of highly aggressive acts were similar
between the sexes. Therefore, despite males having proportionally larger weapons in one species (as predicted by sexual selection),
they fought similarly to females. Our results hint that fighting might not necessarily be the source of selection for sexual dimorphism we
typically expect. Other sources, such as the frequency of fighting and predation pressure, might be selecting larger claws in males
despite the similar fights, while fecundity costs might downsize female claws. We highlight that comparing female with male weapons
and the associated fighting behavior shows that selection on weapons is not as straightforward as we might think.

Significance statement
We show that studying the highly neglected female weapon allometry and usage allows us to infer on the selective process acting on
animal weapons. We show here that males of one species have proportionally larger weapons, and, despite that, males fight similarly to
females in every aspect analyzed. Therefore, fighting per se might not be the sole source of selection to explain sexual dimorphism.
Other sources, such as frequency of fighting, and how expensive it is to produce gametes might be additional sources of selection.

Keywords Animal weapons . Animal contests . Female fighting

Communicated by T. Breithaupt
Electronic supplementary material The online version of this article Introduction
(https://doi.org/10.1007/s00265-019-2741-5) contains supplementary
material, which is available to authorized users.
Animals typically fight over scarce resources (Hardy and
Briffa 2013), and the individual with the highest fighting abil-
* Alexandre V. Palaoro
alexandre.palaoro@gmail.com
ity tends to win the fight and monopolize the resource (Vieira
and Peixoto 2013). Over evolutionary time, the continuous
1
Departamento de Ecologia e Evolução, Universidade Federal de process of overcoming rivals and monopolizing resources
Santa Maria, Santa Maria, Brazil might lead to the evolution of animal weapons: specialized
2
Instituto Federal de Educação, Ciência e Tecnologia morphologies that increase the individual’s ability to over-
Farroupilha-Campus São Vicente do Sul, São Vicente do Sul, Brazil power rivals (McCullough et al. 2016). Weapons have
3
Departamento de Zoologia, Universidade Federal do Rio Grande do attracted considerable attention due to (sometimes) exagger-
Sul, Porto Alegre, Brazil ated proportions and conspicuous behaviors associated with
4
LAGE do Departamento de Ecologia, Universidade de São Paulo, their use (Emlen 2008; McCullough et al. 2016; Rico-Guevara
São Paulo, Brazil and Hurme 2019). However, despite the advances in the last
127 Page 2 of 11
few years (McCullough et al. 2014, 2016; Rico-Guevara and
Hurme 2019), most of that knowledge comes from males. The
few reports on female weapons focus on specific contexts,
such as species with sex-role reversal (e.g., Jacana birds,
Emlen and Wrege 2004). However, males and females can
bear weapons simultaneously, although female weapons in
these instances tend to be smaller than the males’ (Emlen
2008; Hare and Simmons 2018). These small weapons, how-
ever, do not prevent females of crayfish, antelopes, and beetles
from fighting and monopolizing resources similarly to males
(Figler et al. 1999; Stankowich and Caro 2009; Watson and
Simmons 2010). Hence, by not studying females, we are ig-
noring considerable information on sexual selection
(Berglund 2013; Hare and Simmons 2018).
Social selection is typically evoked to explain female
weapons (Tobias et al. 2012; Stockley and Campbell
2013). It states that females fight for resources not directly
related to mating opportunities, such as food or shelter, or that
females use their weapons to fend off predators (sensu Tobias
et al. 2012). If true, then the female phenotypic optimum
would be a weapon that is larger than zero. However, weapons
are theoretically costly to produce and maintain (Allen and
Levinton 2007; O’Brien et al. 2019). The physiological cost
of weapons might, for instance, decrease the number of eggs a
female produces. Thus, despite females having a potential
source of social selection acting on weapon size, fecundity
selection will keep the size of the weapon small, if “typical”
sex roles are maintained (Preziosi and Fairbairn 1997; Reeve
and Fairbairn 1999). Males, on the other hand, also have a
selective pressure for fighting, but they are also typically un-
der sexual selection to monopolize mating partners (Emlen
2008). Thus, fighting would be accompanied by an additional
source of sexual selection acting on weapon size. If fighting
and sexual selection act together in the same direction, and if
males are not under any kind of fecundity selection (such as
paternal care), the male phenotypic optimum is expected to be
higher than the female’s phenotypic optimum. Males should
thus have larger weapons than females. But weapon absolute
size is typically positively correlated to body size without
necessarily indicating sexual selection—larger individuals
should have larger traits (Bonduriansky 2007; Voje 2016). If
weapons are indeed more important for males, theory expects
that males should have proportionally larger weapons than
females. In a log-log regression between weapon size and
body size, then, the slope of the male should be larger than
one (positive allometry sensu Huxley 1924), while the female
slope should be equal or a bit smaller than one (isometry).
That indicates that male weapons receive more resources dur-
ing growth than female weapons, which make them dispro-
portionately larger than female weapons.
Proportionally larger weapons suggest that fighting is also
proportionally more important in that group (e.g., Goyens
et al. 2014, 2016). If fighting is more important in one group,
Behav Ecol Sociobiol (2019) 73: 127
then the resource value is also higher. Resource value is an
important modulator of animal contests (Hardy and Briffa
2013), and the higher it is, the higher the motivation to fight
is (Arnott and Elwood 2009). Higher motivation leads to lon-
ger contests, to individuals being more aggressive, and for
fights to be more intense (Arnott and Elwood 2009). That is
why sexual dimorphism is typically explained by fighting:
more intense selection on weapons should be correlated with
differences in fighting intensity between males and females,
and between species (Lailvaux and Irschick 2007; Charles and
Ord 2012). Hence, it is expected that strong selection on
weapons should have an indirect effect on fighting behavior.
By comparing female weapon morphology and behavior with
male weapon morphology and behavior, we might gain in-
sights on how the strength of sexual selection influences
weapon morphology and fighting behavior.
The freshwater anomurans of the genus Aegla (Decapoda)
offer a unique opportunity to test the impact of female
weapons on sexual selection. Males readily fight in laboratory
conditions, even in the absence of resources (Ayres-Peres et al.
2011; Palaoro et al. 2013). And males were also reported
fighting in the headwater streams they inhabit (M.M.
Dalosto, pers comm. during the experiments of Baumart
et al. 2015). However, there are reports that females fight as
well (Ayres-Peres et al. 2015). Interestingly, males do not fight
for direct access to females before copulation. Instead, males
have shorter and less intense fights when receptive females are
near (Palaoro et al. 2013). But males might fight to ensure that
they are the only individual to mate with a female because
male aeglids defend females after copulation (Almerão et al.
2010). Females, on the other hand, probably fight for food and
shelter. Aeglids are most abundant in pool areas in headwater
streams, areas in which leaf litter accumulates (Bücker et al.
2008; Cogo and Santos 2013; Zimmermann et al. 2016). Since
individuals are abundant in those areas, and that aeglids have a
preference for soft prey rather than leaf litter (Cogo et al.
2018), it is likely that females are fighting for their preferred
food within pool areas. Therefore, male and female aeglids
fight, which ensure that both sexes are under selection for
fighting, but only males may be under sexual selective
pressure.
Our goal is to compare the neglected female weapons and
the associated behavior with the more-studied male weapons
for two species of the genus Aegla. Both sexes fight and both
sexes bear weapons (Bueno et al. 2016), which provides a
unique opportunity to test the predictions of social selection
on female weapons. We hypothesize that females solely under
social selection will bear proportionally smaller weapons
when compared with males—because males are under social
and sexual selection simultaneously (Fig. 1). Furthermore, if
males are also under sexual selection for weapon size, we also
expect male fights to be longer, more intense fights, and for
males to be more aggressive than females.
Behav Ecol Sociobiol (2019) 73: 127
Methods
Morphological data
We used previously obtained data from other studies (Palaoro
et al. 2014 for A. longirostri; Trevisan and Santos 2014 for
A. manuinflata) to compare the morphology of the claw be-
tween males and females. We thus obtained data from 117
adult individuals of A. longirostri (42 males and 75 females)
and 614 adult individuals of A. manuinflata (343 males and
271 females). We obtained measures of carapace length (CL,
measured from the orbital sinus to the posterior edge of the
carapace), length of the left claw propodus (measured from the
articulation with the carpus to the tip of the fixed finger), left
claw height (measured as the broadest part of the claw), length
of the right claw propodus, and right claw height for all
individuals.
With those data, we tested for sexual dimorphism on the
major (left) claw for each species separately. We performed
two ordinary least squares (OLS) regression for each species,
one using the major claw length as the dependent variable and
another using the major claw height as the dependent variable.
We log-transformed CL, the major claw length and height
prior to testing. Log-transforming linearizes the exponential
properties that arise from growth—log approximations allow
us to infer on proportions in a linear scale, rather than an
exponential scale (following Huxley 1924 and Pélabon et al.
2014). We also centered the log-transformed CL to facilitate
comparisons between sexes. As can been seen in Fig. S2,
males and females might differ considerably in the size of their
traits. If data is not centered around the mean, any comparison
that involves body size requires an extension of the fitted line
of one sex to the other sex. Centering thus allows comparisons
of the scaling relationships of both sexes without extending
the fitted lines beyond the range of the data (Schielzeth 2010).
Our independent variables were thus the log-transformed and
mean centered CL, sex, and their interaction. We did not in-
clude species as an independent variable because our goal was
to test for differences in claw allometry between males and
females, adding the species factor would add unwanted com-
plexity and noise in our data. Thus, we performed separated
analyses for both species.
Since selection should also act more strongly on the
major claw than the minor claw (Mariappan et al. 2000),
the degree of heterochely might also hint into the strength
of social/sexual selection on the major claw. Since both
major and minor claws grow under the same environmen-
tal and nutritional conditions, a proportionally larger ma-
jor claw might indicate a proportionally larger allocation
of resources to a strong claw that is mainly used for fight-
ing. Larger heterochely thus means proportionally higher
allocation to strong claws (Fujiwara and Kawai 2016). To
do that, we performed a sexual dimorphism test on the
Page 3 of 11 127
degree of heterochely. We calculated the degree of
heterochely by subtracting the length of the minor claw
from the length of the major claw. Higher values would
indicate that the sex and/or species invested more on the
left claw (the major claw). Then, to test if females and
males differed in their degree of heterochely, we per-
formed a one-way ANOVA: our dependent variable was
the degree of heterochely, while our independent variable
was sex. Once again, we performed separate ANOVAs for
both species. We performed all analyses in the R software
(R Core Team 2018).
Behavioral data
We combined published data from male fights to novel data on
female contest behavior. Data from males came from Ayres-
Peres et al. (2015), which analyzed the contest behavior of
A. longirostri (N = 9 pairs) and A. manuinflata (N = 8 pairs).
For the female contests, we sampled adult females of
A. longirostri (N = 10 pairs) in a tributary of the Vacacaí-Mirim
River, Jacuí River Basin, Santa Maria, Brazil (29°40′13″ S;
54°45′44″ W) and adult females of A. manuinflata (N = 10 pairs)
in the Passo de Taquara Stream, Uruguay River Basin, São Pedro
do Sul, Brazil (29°36′ 01″ S; 54°10′37″ W). We used only
intermolt adults with all appendages intact in the experiments
(Colpo et al. 2005; Trevisan and Santos 2014).
We brought the females to the laboratory where they accli-
mated for at least 7 days prior to the experiments to control for
the effects of any previous social interactions (Moore 2007). We
fed the crabs every other day with fish fillet to standardize hunger
motivations, with the last feeding occurring 48 h prior to the
experiments. We maintained the crabs in individual aquaria
(2 L), which were visually and chemically isolated from each
other, in a room with controlled temperature (17.5 ± 1.7 °C) and
a 12D:12N photoperiod. We paired females according to their
CL, with a maximum variation of 1 mm between individuals of
the pair (Palaoro and Briffa 2017).
For the experiments, we used a similar aquarium to the one
from the published experiment (Ayres-Peres et al. 2015): a 20 ×
12 × 13 cm aquarium separated in three equal compartments by
removable opaque plastic dividers. One member of each pair was
simultaneously placed in the opposite side of the aquarium and
allowed to acclimate to the aquarium for 20 min. Afterwards, we
removed the dividers and allowed the females to interact for
another 20 min. We filmed the interaction with a Sony®
Handycam DCR-DVD 108 video camera positioned perpendic-
ularly 26 cm above the aquarium. We always performed the
experiments at night, as aeglids are nocturnal animals, and day-
light might influence their behavior. We also used red incandes-
cent light bulbs (40 W) to illuminate the aquarium because crus-
taceans have low sensitivity to this wavelength (e.g., Dalosto and
Santos 2011). We chose an experimental design without re-
sources to allow for a direct comparison between males and
127 Page 4 of 11 Behav Ecol Sociobiol (2019) 73: 127

females, and because the presence of sexually receptive females
decreases male aggression (Palaoro et al. 2013). After the inter-
action, we returned the females to their individual aquaria, where
they remained for at least a week before they were released back
in their collection sites.
As mentioned earlier, we compared female contests dynamics
with previously published data on male contest dynamics
(A. longirostri and A. manuinflata; Ayres-Peres et al. 2015). We
controlled for as many factors as possible to increase the compa-
rability of our data to that of the previous study, such as
performing the experiments in the same temperature, using the
same protocols, and sampling on the same locations. We calcu-
lated contest duration similarly to Ayres-Peres et al. (2015): we
summed the duration of the first three aggressive bouts, which
are considered the most informative. A bout began when one
individual approached to within one body length of its opponent
and the opponent did not flee. We deemed the bout finished when
one individual retreated and did not approach the other for at least
5 s. In each bout, the individual that retreated was deemed the
loser of the bout, and its opponent the winner of the bout. The
individual that won most bouts was deemed the winner of the
interaction, and its opponent the loser. In all cases, an observer
that was blind to the crab’s identity, sex, and species, and un-
aware of the study’s hypotheses, analyzed the recordings.
For each interaction, we quantified the total time spent
fighting (sum of the duration of the three bouts), the pro-
portion of time spent by winners on highly aggressive be-
haviors (time spent on unrestrained use of the claws rela-
tive to total time spent fighting, see (Ayres-Peres et al.
2011)), and the latency to start the contest (time spent until
the first bout began). To test for differences in fight dura-
tion, we used a generalized linear model (GLM) with a
Gamma error structure and a log link, with sex as the pre-
dictor variable. For the proportion of time spent on highly
aggressive behaviors, we used a beta regression, in which
the response variable was the proportion on highly aggres-
sive behaviors, and the predictor variable was sex. Lastly,
we log-transformed the latency to start a contest to use it as
a response variable in an ordinary least squares regression,
while our predictor variable was sex. We performed an
analysis for the species separately. And all analyses were
performed in R software using “nlme” package for GLM

2.9 3.2
a b c
2.0
2.8
3.0

Claw length (log)

2.7
1.5
Intercept

2.8
Slope

2.6

1.0 2.6
2.5

2.4 2.4
0.5
2.3 2.2
Female Male Female Male −0.3 −0.2 −0.1 0.0 0.1 0.2 0.3
sex Sex Cephalothorax length (log−centered)

d e f
2.8 1.4
3.0
1.2
Claw length (log)

2.6
Intercept

Slope

1.0
2.5
2.4
0.8

2.2
0.6 2.0

2.0 0.4
Female Male Female Male −0.2 −0.1 0.0 0.1 0.2 0.3
Sex Sex Cephalothorax length (log−centered)

Fig. 1 Females (light gray) have shorter claws than males (dark gray) on
average (intercept value, a and d), while only males of Aegla longirostri
(top row) have proportionally longer claws than females (b). Males of
A. manuinflata (bottom row) have proportionally similar claws to females
(e). Panel a shows the intercepts (dots) and 95% confidence intervals
(whiskers) of males and females of A. longirostri, and panel d shows
the same information for A. manuinflata. Panel b shows the slopes and
the corresponding 95% confidence intervals for A. longirostri, and panel e
for A. manuinflata. To obtain these values, we used an ordinary least
squares regression of log-transformed claw length as a function of cen-
tered and log-transformed cephalothorax length. In panels c and f, we
show males (dark gray) and females (light gray) of A. longirostri and
A. manuinflata, respectively. Dashed line indicates the regression model
for females and gray solid lines for males
Behav Ecol Sociobiol (2019) 73: 127 Page 5 of 11 127

analyses and “betareg” package for the beta regression
(Cribari-Neto and Zeileis 2010; Pinheiro et al. 2018). All
figures and graphics were also made using the R software.
Results
Claw allometry
Females had, on average, smaller claws regarding length and
height regardless of the species (i.e., lower intercepts, Figs.
1a, d and 2a, d). However, only A. longirostri males had propor-
tionally larger claws regarding length and height (steeper slopes,
Figs. 1b and 2b). Males A. manuinflata had proportionally sim-
ilar claws regarding length and height (shallower slopes, Figs. 1e
and 2e). Fitted regression lines for each species can be seen in
Fig. 1c, f and Fig. 2c, f. Statistical results can be seen in Table 1.
Finally, females were also less heterochelic than males of both
species (one-way ANOVA, A. longirostri, F1,79 = 32.63,
P < 0.0001, Fig. 3a; A. manuinflata, F1,272 = 326.8, P < 0.0001,
Fig. 3b).
Contest dynamics
One pair of females of each species did not fight. We
removed those female pairs from the analyses, but results
were similar (data not shown, but available on GitHub).
Thus, we present the results with all females in the anal-
yses. Females fought similarly to males, both qualitatively
and quantitatively. Females began contests by ap-
proaching each other face to face and whipping their an-
tennae on the rival. Contests then proceeded to physical
contact, in which females touched each other’s claws
without trying to grab the rival. Lastly, females adopted
the same “fighting stance” of male aeglids (Palaoro et al.
2014), in which individuals bend their smaller claw to-
wards the substrate keeping the larger claw open towards
the rival while trying to grab the rivals’ smaller claw.
Regarding contest quantitative variables, contest dura-
tion was similar regardless of sex (Table 2, Fig. 4a, d).
Females also spent a similar proportion of time on highly
aggressive behaviors (Table 2, Fig. 4b, e) and showed a
similar latency to start a contest (Table 2, Fig. 4c, f).

a 2.5 b c
2.4 2.6

2.0 2.4

Claw height (log)

2.2
Intercept

Slope

1.5 2.2

2.0 2.0
1.0

1.8
1.8
0.5
1.6
Female Male Female Male −0.3 −0.2 −0.1 0.0 0.1 0.2 0.3
sex Sex Cephalothorax length (log−centered)

1.8
d e f
2.4
2.5
1.6
2.2
Claw height (log)
Intercept

1.4 2.0
Slope

2.0

1.2
1.8
1.5

1.6 1.0

1.0
1.4 0.8
Female Male Female Male −0.2 −0.1 0.0 0.1 0.2 0.3
Sex Sex Cephalothorax length (log−centered)

Fig. 2 Females (light gray) have claws with less height than males (dark
gray) on average (intercept value, a and d), while only males of Aegla
longirostri (top row) have proportionally larger claw height than females,
although larger on average (b). Males of A. manuinflata (bottom row)
have proportionally similar claws to females (e). Panel a shows the
intercepts (dots) and 95% confidence intervals (whiskers) of males and
females of A. longirostri, and panel d shows the same information for
A. manuinflata. Panel b shows the slopes and the corresponding 95%
confidence intervals for A. longirostri, and panel e for A. manuinflata.
To obtain these values, we used an ordinary least squares regression of
log-transformed claw height as a function of centered and log-
transformed cephalothorax length. In panels c and f, we show males (dark
gray) and females (light gray) of A. longirostri and A. manuinflata, re-
spectively. Dashed line indicates the regression model for females and
gray solid lines for males
127 Page 6 of 11 Behav Ecol Sociobiol (2019) 73: 127

Table 1 Results of ordinary least squares regressions performed for Discussion

Aegla longirostri (a and b) and Aegla manuinflata (c and d) separately.
We present the results for regression using claw length (a and c) or claw
height (b and d) as response variables, and body size (i.e., cephalothorax Our goal was to bring light to female weapons by studying its
length), sex (male or female), and their interaction as predictor variables. allometry and the corresponding contest behavior while com-
Italic indicates significant factors. Df degrees of freedom paring with the more studied male weapons and behavior. We
Aegla longirostri found that males have, on average, larger claws than females.
(a) Claw length Sum of squares Df F P But, only in one species do males have proportionally larger
Cephalothorax length 2.164 1 224.61 < 0.001
claws than females. It thus seems that predictions regarding
Sex 3.423 1 355.24 < 0.001
selection on weapon allometry are not as straightforward as
Interaction 0.118 1 12.127 < 0.001
expected when we consider the allometry of female weapons.
Furthermore, males also do not fight differently from
Residuals 0.742 77 – –
females—neither qualitatively nor quantitatively. Therefore,
(b) Claw height Sum of squares Df F P
not only are the predictions regarding weapon allometry
Cephalothorax length 2.81 1 219.35 < 0.001
straightforward, but they are also not correlated to contest
Sex 4.69 1 366.36 < 0.001
behavior. Below, we discuss potential explanations to why
Interaction 0.13 1 10.39 < 0.001
males and females do not differ as much as predicted and
Residuals 0.986 77 – –
summarize what we learned from female weapons.
Aegla manuinflata Fighting is one of the forefront explanations for sexual
(c) Claw length Sum of squares Df F P
dimorphism in weapons. It is evoked to explain the marked
Cephalothorax length 2.96 1 424.69 < 0.001
sexual dimorphism in the claws not only of aeglids, but most
Sex 36.05 1 5168.60 < 0.001
Decapoda (e.g., swimming crabs: Pinheiro and Fransozo
1993; crayfish: Vlach and Valdmanová 2015; aeglids: Adam
Interaction 0.01 1 0.767 0.382
et al. 2018; porcelain crabs: Baeza and Asorey 2012; shrimps:
Residuals 1.88 270 – –
Baeza et al. 2010; snapping shrimps: Hughes et al. 2014), and
(d) Claw height Sum of squares Df F P
several other taxa (reviewed in Hare and Simmons 2018). A
Cephalothorax length 4.32 1 498.54 < 0.001
common expectation, thus, is that females fight less than bet-
Sex 54.41 1 6283.63 < 0.001
ter-armed, large males. However, since females fought simi-
Interaction 0.01 1 1.13 0.288
larly to males, we cannot expect the common Darwinian sce-
Residuals 2.34 270 –
nario (large weapons increase access to females) to hold in this
case; especially when we consider that male A. longirostri
have shorter and less intense fights when sexually receptive
females are present (Palaoro et al. 2013). Thus, other selective
pressures need to be considered to explain why males and
females differ in their weapons, while not differing in fighting.

6
a b
Difference between left and right claw

Difference between left and right claw

10
5

8
4

6
3

4
2

1 2

0 0

Female Male Female Male

Sex Sex

Fig. 3 Females (light gray) are less heterochelic than males (dark gray), quartiles, and the whiskers the minimum and maximum. Gray dots
regardless of the species. The y-axis represents the index of heterochely, represent each individual. Aegla longirostri is represented on (a) and
which is a subtraction between the left claw length and the right claw Aegla manuinflata on (b)
length. Horizontal line represents the median, the box the first and third
Behav Ecol Sociobiol (2019) 73: 127 Page 7 of 11 127

Table 2 Results of the tests performed for Aegla longirostri and Aegla fighting between males is frequently seen during austral
manuinflata regarding sex differences in fight duration (a), frequency of
spring nights in the field (M. M. Dalosto, pers. comm., Fig.
highly aggressive behaviors (b), and latency to start a contests (c). We
used a generalized linear gamma regression in (a), a beta regression in (b), S1.). If the frequency of fighting dictates how a weapon is
and an ordinary least squares regression in (c). We did not find shaped, then we should not expect the sexes to differ in how
significance in any of the tests. Df degrees of freedom they fight—only in how often they fight. Furthermore, males
Aegla longirostri might also need larger weapons to maintain stable hierarchies.
(a) Fight duration χ2 Df F P Although hierarchy formation has not been studied in aeglids,
Sex 1.81 1,17 – 0.192
evidence from crayfish strongly suggests that aeglids also
(b) Aggressive behaviors χ2 Df F P
form and maintain hierarchies (Dalosto and Palaoro in
Sex 0.044 1,17 – 0.833
press). If so, then it is possible that males use their strong
claws to fight for, or to maintain, a high social status that could
(c) Latency Sum of squares Df F P
increase their indirect access to mates (Moore 2007).
Sex 0.497 1,17 0.366 0.553
We also cannot discard the impact of female preference and
Aegla manuinflata genetic correlations on weapon size. First, female preference
(a) Fight duration χ2 Df F P for large, more conspicuous male weapons might increase
Sex 0.479 1,16 – 0.305 male weapon size (Siqueira et al. 2013). Given that both sexes
(b) Aggressive behaviors χ2 Df F P fight similarly, under a Fisherian selection scenario (i.e.,
Sex 1.393 1,16 – 0.237 runaway and/or sexy son, Andersson 1994), the optimum
(c) Latency Sum of squares Df F P weapon size would be similar for both males and females—
Sex 0.408 1,16 0.722 0.408 but female preference would push male weaponry away from
the female optimum and into exaggerated sizes. However, the
nocturnal habits and small home ranges of aeglids (Baumart
et al. 2015), along with the female’s more cryptic behavior (as
Females might bear smaller weapons because of the higher
seen by their low capture rate in traps, Dalosto et al. 2014),
energetic constraints imposed by large weapons when compared
decrease the likelihood of females sampling lots of males,
with males. Carrying eggs and caring for them until they are
which in turn might decrease the intensity of sexual selection
released as juveniles, as is the case of aeglids, is energetically
by female preference (Muniz and Machado 2018). Second,
costly (López-Greco et al. 2004). And developing and maintain-
female weapons can occur because of genetic correlations
ing a weapon is also costly (Levinton and Allen 2005; O’Brien
between males and females (Berglund 2013; Hare and
et al. 2019). Given a finite pool of resources, females might have
Simmons 2018). If weapon expression is tied to nonsexual
a higher fitness payoff by investing on more eggs than on large
genes, then male and female weapon might also be tied: the
weapons, while males might be the opposite, investing more on
male optimum would pull the female optimum together de-
weapons than in sperm. If true, then any investment made in
spite any cost associated (Bonduriansky and Chenoweth
weapons by females would decrease their fitness by decreasing
2009). Although genetic correlation explains why females
the number of eggs, for instance. Our results of heterochely in-
might bear weapons (and exaggerated weapons as well), it
dicate that, although males haver larger claws, females also invest
does not explain why females are fighting similarly to males.
in fighting claws. But, the proportionally larger abdomen in fe-
Interestingly, sexual dimorphism in claw size (regardless of
male aeglids evidences the selection for fecundity in females
the measure) differed between species. Aegla longirostri
(Barría et al. 2014). Therefore, regardless of the importance of
males had proportionally larger claws than females, while
fighting, fecundity selection could hamper females from increas-
A. manuinflata males only had larger claws on average. We
ing their weapon size, generating the sexual dimorphism we
refrain from making any comparisons between species be-
observe on weapons.
cause that is not our goal and because we do not have enough
Another explanation resides not in contest dynamics per se,
information on behavior and ecology for both species. A
but rather in the frequency in which individuals of different
broader comparative analysis of claw morphology would pro-
sexes engage in contests. By engaging in contests more fre-
vide more insights on the evolution of male and female
quently, fighting might be more important in one sex than the
weapons than we can achieve with our sample. However,
other. For instance, species of stalk-eyed flied (i.e., Diopsidae)
these differences do highlight our need for more information
that fight more are more dimorphic regarding their eyestalks
on closely related species. Two recent meta-analyses on fight-
than species that do not fight (Rohner and Blanckenhorn
ing behavior (Vieira and Peixoto 2013; Pinto et al. 2019)
2018). In aeglids, males are found in higher densities within
showed that the bulk of our knowledge comes from single
streams during the peak of the breeding season. Thus, it is not
species—studies on closely related species are rare. Our abil-
unlikely that males encounter each other more frequently than
ity to make inferences is thus hindered by the lack of informa-
females do (Dalosto et al. 2014; Baumart et al. 2015). Indeed,
tion on how congenerics and females behave.
127 Page 8 of 11 Behav Ecol Sociobiol (2019) 73: 127

Proportion of time spent in highly aggressive behaviors

a 1.0 b c
7
1000

Latency to start contest (log)

0.8 6
800
Fight duration (s)

0.6 5
600

400 0.4 4

200 0.2 3

0 0.0 2

Female Male Female Male Female Male

Sex Sex Sex
Proportion of time spent in highly aggressive behaviors

350 d 1.0 e f
7.0
300

Latency to start contest (log)

0.8 6.5
250
Fight duration (s)

6.0
200 0.6
5.5
150
0.4
5.0
100
0.2 4.5
50

0 4.0
0.0

Female Male Female Male Female Male

Sex Sex Sex

Fig. 4 Females (light gray) fought similarly to males (dark gray) to start a contest (c, f) did not differ between the sexes. Horizontal line
regardless of the variable measured and species (Aegla longirostri on represents the median, the box the first and third quartiles, and the
top row and Aegla manuinflata on bottom row). Fight duration (a, d), whiskers the minimum and maximum. Gray dots represent each
proportion of time spent in highly aggressive behaviors (b, e), and latency individual. Gray dots represent each fighting pair

This can be better understood if we look for the possible females are added, we see that despite high behavioral simi-
effects of natural selection in animal weapons, such as when larity, weapons might not be under the same selective pres-
predation differs between sexes or species (Stankowich 2012; sures. For instance, males and females might be under differ-
Kojima et al. 2014). In crayfish, for example, although the ent predation pressures because their escape responses differ
base pattern of agonistic behavior is shared among most spe- when females are gravid (Benso-Lopes et al. 2019).
cies (see Moore 2007), different predation pressures correlate Therefore, more information on females might allow us to
to differences in claw allometry. Such is the case of Faxonius make stronger inferences on the evolutionary processes and
crayfish that bear large claws and are mainly preyed upon by patterns behind both male and female weaponry.
fish. However, Faxonius immunis is mostly preyed upon by Overall, studying female weapons provided insights into
mammals and birds, against which large claws are ineffective. the evolution of animal weapons. The Darwinian view of sex-
Thus, while most Faxonius species preyed by fish show large ual selection implies that males are selected for large weapons
claws, Faxonius immunis has smaller claws possibly because because large weapons would increase male fitness
of a different predation pressure (Garvey et al. 1994; Tierney (Andersson 1994). However, we showed here that females
et al. 2000). In aeglids, recent work has shown a high behav- can also bear—and use—weapons in fights. And despite the
ioral similarity between males of different species (Ayres- males of one species having proportionally larger claws, fe-
Peres et al. 2015). Since most males bear larger claws than males fight similarly to males. Thus, social/sexual selection
females (Bueno et al. 2016), it is logical to interpret that males might not be as straightforward as we consider. We proposed
fight more than females. However, when our results for four nonexclusive explanations for why male and female
Behav Ecol Sociobiol (2019) 73: 127
weapons are dimorphic while fighting similarly. The main
mechanisms are stronger fecundity selection on females and
higher frequency of contact fighting in males. Future studies
could investigate the magnitude of the selection imposed by
how frequently animals fight. Furthermore, we also need to
understand better the costs for females to bear weapons. Are
female weapons hampered by fecundity costs? A third venue
for further research is the impact of natural selection on male
and female weapons. Female bovids use their horns against
predators (Stankowich and Caro 2009), and decapods, as stat-
ed above, do the same. Thus, we need to understand the im-
pacts of predation on selecting male and female weaponry.
Here, by studying female weapons and associated behavior,
we have shown that sexual dimorphism might not be ex-
plained entirely by fighting. And that selection on animal
weapons might not as straightforward as we think.
Acknowledgments We thank Glauco Machado for the fruitful discus-
sions on female weapons.
Data accessibility All code and data are deposited on GitHub: https://
github.com/alexandrepalaoro/femaleweapons.
Funding information MMD thanks CAPES (process no: 23081.048599/
2018-42) and AVP thanks FAPESP (process: 2016/22679-3) for the post-
doctoral grants. SS thanks CNPq for the productivity grant (process:
311142/2014-1).
Compliance with ethical standards
Ethical approval All applicable international, national, and/or institu-
tional guidelines for the care and use of animals were followed.
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