Absence of handedness in female fiddler crabs, Uca tangeri,

and the effect of left claw usage on feeding rate

ABSTRACT
Fiddler crabs use their claws for foraging. Unlike male crabs, which possesses a non-
functioning enlarged claw, female crabs have two functioning claws that they can use
while foraging. This raises the question of whether female crabs exhibit a preferred
handedness during foraging. We investigated the presence of preferred handedness in
female crabs and its potential correlation with foraging effectiveness. We examined 18
short videos capturing the foraging behavior of female fiddler crabs and analyzed the
data. Here we show that female fiddler crabs do not exhibit a significant preferred
handedness during foraging. This aligns with our initial hypothesis that the absence of
preferred handedness is favorable for optimal foraging. However, our results did reveal
that there is a positive correlation between the utilization of the left claw and a higher
feeding rate among the female fiddler crabs.
This suggests that preferred handedness does indeed influence the foraging
effectiveness of female fiddler crab.

Key words
Fiddler crabs, female, preferred handedness, foraging, feeding rate.

Introduction
Fiddler crabs are foragers that mostly eat algae, bacteria, and fungi that grow in the
mudflats, using their claws to transfer the food to their mouths. Female fiddler crabs use
both claws to forage for food, either alternating or using both claws at the same time. If
a female uses her right or left claw more to feed, we consider her right- or left-handed.
Handedness involves sensory and motor skills (Michel et al., 2018). It is an important
trait that is associated with the imbalances between the left and the right hemisphere in
the motor hand region, specifically after the enteral sulcus’s middle section. These
imbalances manifest on the opposite side of the brain from where the hand movement
started (Margiotoudi et al., 2019).
Male fiddler crabs are known for possessing a significantly larger claw used for
attracting females. We consider their smaller claw as their preferred claw for feeding.
Males in some species of crab have a clear preference, while other species have an
equal distribution of left and righthanded crabs (Rozaimi et al., 2023). However, it is not
known whether female fiddler crabs exhibit a preference like the males do. There is
evidence of males and females in some species either having a preferred claw to feed
(Pescinelli et al., 2020), while in others there is no preference to be found at all (Xiang
et al., 2020).
Prior studies have shown that in most species of fiddler crabs, the males show an
evenly distributed occurrence of an enlarged left or right claw (Backwell et al., 2007).
Other studies have also shown that Uca Chlorophthalmus female fiddler crabs can feed
50% faster than males (Weis and Weis 2004).
Our aim of this research paper is to explore and analyse the preferred handedness of
female fiddler crabs and whether this influences their feeding rate.
We hypothesize that the feeding rate of female fiddler crabs will be influenced by the
presence of a preferred claw. If a female fiddler crab lacks a preferred claw, she can use
both claws more efficiently compared to another who uses one claw more frequently.

Methods
Our study focused on a population of fiddler crabs, Uca tangeri, living in the Royal
Burger’s Zoo in Arnhem. We collected data from 18 recordings of female crabs with two
functioning claws, each video lasting two minutes, to examine the frequency at which
they used their left claw and/or right claw. We examined the frequency of the use of the
claws by counting each time the crabs successfully transferred food to their mouths, for
both claws (n1= 36). We conducted an analysis to determine the ratio of preferred
handedness for female crabs and whether it correlates to their feeding rate.
The most significant approach to determine the ratio of preferred handedness involves
multiple key steps. Firstly, employing a paired t-test, we assessed whether an overall
significant difference existed in the frequency of left or right claw used among all female
fiddler crabs. Next, we conducted individual binomial tests on each female crab to
explore potential preferences for utilized claw when foraging. Lastly, we conducted a
Pearson’s correlation test to examine whether the frequency of used right or left claw
correlated to the feeding rate of the female fiddler crabs.

Results
The female crabs showed no significant difference in overall preferred handedness
between them (t17 = -1.06, p = 0.31), however three females (3, 11, and 13) showed a
left claw preference with p<0.05 while the rest showed no preference (Table 1). Use of
left claw and feeding rate was positively correlated, r(16) = 0.49, p = 0.04 (Fig 1).

Table 1. Binomial test results for right and left claw preference, expected 0.5.
Right Claw Left Claw
Observation P-value P-value
1 0.84 0.16
2 0.71 0.29
3 0.98 0.02
4 0.29 0.71
5 0.47 0.53
6 0.31 0.69
7 0.60 0.40
8 0.67 0.33
9 0.77 0.23
10 0.25 0.75
11 0.99 0.01
12 0.53 0.47
13 0.99 0.01
14 0.11 0.89
15 0.10 0.90
16 0.52 0.48
17 0.33 0.67
18 0.70 0.30
Fig. 1. Correlation between left claw usage and feeding rate. Relative left claw usage calculated by
subtracting use of right claw from use of left claw, therefore females who used their right claw more lie to
the left of the y-axis.

Discussion
Our findings suggest that female fiddler crabs do not exhibit a clear preference for
handedness, as both claws were frequently used for feeding activities. However, we did
discover a correlation between the claw utilized by female fiddler crabs during feeding
and the rate at which they are feeding. Specifically, females that primarily used their left
claw demonstrated higher feeding rates.
Female crabs show no claw preference when feeding, which aligns with the absence of
any apparent biological benefit in using one specific claw (Swanson et. Al, 2013).
Female crabs do have the benefit that both claws can be used for feeding.
Even though there was no significant preference for the right or left claw, a small skew
in the claw used for feeding was evident. Additionally, a correlation between the feeding
rate and the preferred claw used more during feeding was established, even though
there is no known biological advantage associated with this behavior. Also, in research
about other crab species, which also possess two functional claws, no correlation
between feeding behavior and handedness has been found (Wang et al., 2021).
The difference in feeding rate observed may be explained by some unaccounted
factors. The size of the ingested particles influences the feeding rate of fiddler crabs
(Robertson et al., 1982), and the size of the crabs themselves was unknown and could
also influence the feeding rate (Valiela et al., 1974). Further research should investigate
these factors to explain the correlation between the feeding rate and the claw used most
for feeding by female fiddler crabs.

Author statement
Danica Cunningham: data collection, methods, results. Hanna Vargas: data
collection, introduction, methods, editing, reference list. Kayleigh Maduro: data
collection, introduction, hypothesis. Luuk Verkerk: data collection, introduction.
Mitchell van Eeden: data collection, abstract, introduction. Teun de Vries: data
collection, discussion.
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