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Comparison of individual and groupbehaviour of varying sizes of the light-blue soldier crabs (Mictyrislongicarpus) in the presence of a predator
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Scott Pegg 2016
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Abstract | |
The aim of this study was
to determine the behaviour of the light blue soldier crab Mictyris
longicarpus (Latreille 1806) in the presence of a simulated predator. The
behaviour of scurrying distance and burying time was assessed with respect to
the crab size and a comparison between individuals and groups. A linear
relationship was found between crab size and scurrying distance with 1
millimetre of crab increasing the distance by 13.01cm (R2=42.34%). The
relationship between crab size and burying time increased at a rate of 0.85
seconds per 1 millimetre of crab (R2=30.98). Comparing individuals
with groups of 5 crabs displayed no significant difference in scurrying
distance, however a significant difference was discovered in burying time. This
is likely to be as M. longicarpus finds safety in numbers, similar to
many other species in the animal kingdom. Behaviour such as this may benefit
the individual but not the group.
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Introduction | |
The light-blue soldier
crab, Mictyris longicarpus (Latreille 1806) is decapod in the mictyridae
family. When fully grown M. longicarpus
had a carapace length of 2.5cm (Rossi &
Chapman, 2003). M. Longicarpus is found from the tropics of New
Caledonia and Singapore all the way south to the temperate regions of southern
New South Wales and Western Australia (Rossi
& Chapman, 2003; Webb &
Bradley, 2004). The crab commonly inhabits the intertidal regions of
bays and estuaries, on flats of sand and muddy sediments (Rossi
& Chapman, 2003; Webb &
Bradley, 2004). During the high tide soldier crabs bury themselves in
the sediment to depths of 300mm (Maitland
& Maitland, 1992) but are usually found in the top 200mm of
sediment (Kelemec, 1979). Under
the sediment they retain an air chamber as they predominantly breathe air (Maitland
& Maitland, 1992). M. longicarpus emerge to the surface on
the ebbing tide (Kelemec, 1979),
however Kelemec (1979) found that crabs may not emerge if the temperature was
to cold. Supporting this Cameron (1966) discovered that in Moreton Bay,
Queensland, soldier crabs were more active on ‘warm sunny days’ rather than on
‘cold overcast days’. The crabs can remain on the flats for over 4 hours (Webb
& Bradley, 2004). During this time they regularly aggregate in
swarms (Cameron, 1966; Rossi
& Chapman, 2003) and can travel up to 450m, feeding for two or more
hours (Cameron, 1966; Webb
& Bradley, 2004). Unlike many other brachyuran species, M.
longicarpus does not reside in one permanent burrow (Dittmann,
1993), instead it can rapidly bury itself with its unique corkscrew
motion (Morton & J.E., 1983),
likely to avoid predation (Takeda, et
al., 1996).
Many studies have been conducted
examining both the physiological aspects and environment factors affecting
soldier crabs behaviour. For example physiological investigations have been
conducted on appendages and locomotion (Sleinis
& Silvey, 1980), visual ability (Kraus
& Tautz, 1981), feeding ability and mechanics (Quinn,
1980; Takeda & Murai, 2004),
respiratory system (Maitland &
Maitland, 1992) and chitosan exoskeleton membranes (Chen,
et al., 2006). Furthermore environmental factors explored include the
effects of varying sand temperature (Kelemec,
1979), salinity concentrations and tolerances (Barnes,
1967), favouring sediment types and grain sizes (Rossi
& Chapman, 2003) and the crabs importance for sediment irrigation
and nitrogen fluxes (Webb & Bradley,
2004). Interspecies interactions investigate mating (Nakasone
& Akamine, 1981 ), population densities (Shih,
1995) and soldier crab swarm behaviour (Cameron,
1966; Unno & Semeniuk, 2011)
where Murakami and team (2014) discovered that soldier crabs behaviour changed
with an increase number of individuals in a group. Whilst intraspecies
interactions have predominantly been studied for food gathering and prey items (Quinn,
1986; Dittmann, 1993; Takagi,
et al., 2010) and food-web and trophic connections (Lee,
et al., 2011; Abdullah & Lee,
2016), it was revealed by Takeda (2010) that fiddler crabs prey on
soldier crabs, preferable of a smaller size, resulting in habitat partitioning
between the two species. Soldier crabs are have also been evidently preyed upon
by gastropods (Huelsken, 2011), various
fish species (Queensland Museum, 1998),
elasmobranchs (Pearce, 2008) and
shoreline birds (Queensland Museum, 1998).
As suggested by Takeda and
team (1996) the crabs would bury to avoid predation. The behaviour of soldier
crabs was also found to change depending on the number of individuals within a
group (Murakami, et al., 2014).
The size of a crab also changed their behaviour in predator avoidance (Takeda,
2010). Acknowledging this, this study investigates the behaviour of
the light-blue soldier crab, Mictyris longicarpus, when the
threat of a potential predator is placed on them. Individual crabs and groups
were investigated to determine whether crab size and group size effected the
burying time and scurrying distance of a crab.
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Materials and Methods |
Study Site | |
The study was conducted at Dunwich, North
Stradbroke Island, Queensland (27⁰29’45”S, 153⁰24’2”E) (Figure 1) on the 14th
and 15th of May, 2016. On the shoreline of Moreton Bay, the tidal
flats of Dunwich are of sandy to muddy sediments (Kruck,
et al., 2009). This area is a semi-diurnal tidal zone, with an average
tidal range of 1.1m during the neap tides (half-moon) and 1.8m during spring
tides (full or new moon) (Morton, et al.,
1987). Both days field work was conducted there was sunny weather,
temperatures 15 to 25⁰C, barometer was between 1020 and 1024 mbar and winds
from the south west direction at less than 10 knots. The moon was at its first
quarter on the 14th of May. Low tides occurred at 1036 (with a
height of 0.67m) and 2210 (0.89m) with high tides at 0322 (2.20m) and 1624
(1.75m) on the 14th May. The 15th May had low tides at
1132 (0.71m) and 2322 (0.86m) and high tides at 0426 (2.05m) and 1729 (1.85m).
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Figure 1 |
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Crab Collection and size | |
Crabs were collected on the sand flats between
2 hours either side of the low tides. Crab size was the distances between the
third pair of appendages (Figure 2). The crabs were kept with other crabs of
similar sizes in buckets of 40cm diameter. The buckets had moist sand and crabs
were kept in the shade until tested. At any time the crabs had a maximum of 30
crabs per bucket. All crabs were released and all examination was complete on
the day. Crabs were separated into three groups; small (3.0-8.0mm), medium
(8.1- 14.0mm) and large (>14.1mm).
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Figure 2 |
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Experimental Design | |
A pen of 5.5m x 1.0m was set up on the
intertidal edge of the sand flat (Figure 3). A crab pen was set up so soldier
crabs could not backtrack on their paths. Crabs were selected as either
individuals or groups of 5 and were placed on a 0.5m square piece of wood
covered by shelter of 15cm x 15cm. After a stabilising period of 30 seconds, a
person walked to the pen and removed the cup to simulate a predator. Timing
started once the crabs walked off the wood and was stopped when crabs commenced
burying. For groups of 5, burying time started once 3 crabs had left the wood
and stopped when 3 crabs had commenced burying. Scurrying distance was measured
by the straight line distance from the front centre of the wood to where the
crab had buried itself. In groups of 5 the group distance was determined as the
average of the 3 median soldier crab distances. If crabs walked in the side of
the pen their trial was not counted.
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Figure 3 |
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Statistical Analysis | |
Individual crab results were categorized into
size categories as to match with the group data. ANOVA tests were conducted
between groups of different crab sizes for both burying time and scurrying
distance.
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Results |
Burying time | |
Overall 88 individual
crabs and 75 crabs in groups were tested. Individual crabs were both analysed
as individual and in size categories to be compared with group crab data. When
separated, individual crabs had 42, 22 and 24 in the small, medium and large
crab categories respectively. Individual sizes of M. longicarpus ranged from 3.0mm to 20.3mm. For the groups, each
category had 5 groups of crabs in them.
Within individual crabs,
burying time varied from 0.9 seconds to 36.7 seconds. A linear relationship was
placed between the crab size and burying time (Figure 4). It was seen that with
one millimetre of size, crab burying time increases by 0.85 seconds. The linear
relationship explained 30.98% of variance within the data (R2=30.98)
(Figure 4). Crab burying time was not significantly different between
individual crabs and groups of crabs for each size category and an overall
comparison (Figure 5). Data had large standard deviations away from the mean
(Figure 5). Groups had burying times from 7-22 seconds in the small category,
9-33 seconds in the medium category and 14-32 seconds in the large category.
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Figure 4 |
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Figure 5 |
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Scurrying Distance | |
Scurrying distance of individual crabs had a
range of 1cm to 430cm. A linear relationship explaining 42.34% of the data (R2=42.34%)
exhibited that with every one millimetre of crab length, M. longicarpus
s are likely to travel an extra 13.01cm (Figure 6). Groups of crabs were found
to not differ significantly to the scurrying distance of individual crabs
(Figure 7). Data had large standard deviations (Figure 7). Group
means varied from 60-194.3cm in the small category, 51.6-203cm in the medium
category and 79.6-282cm in the large category.
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Figure 6 |
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Figure 7 |
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Discussion |
Crab Size Behaviour | |
With warm sunny conditions, the M.
longicarpus were at a peak activity level (Cameron,
1966). Burying time and scurrying distance was found to increase with
the size of crab (Figure 4,6). The ability to burrow quickly is likely to avoid
predation (Takeda, et al., 1996).
Knowing this and considering that crabs of larger sizes have the confidence to
take longer to bury and scurry further it can be suggested that crab size
effects the rate of predation. Smaller crabs are more vulnerable to a larger
group of predators. For example a past study found that soldier crabs of
smaller sizes are more susceptible to predation by fiddler crabs (Takeda,
2010). Larger crabs also had the ability to scurry at a faster pace
than the smaller crabs, hence are more likely to escape a predator without
having to bury. Behavioural adaptations exhibited by soldier crabs include
crabs raising themselves up to appear larger (Takeda,
2010). This is likely to be as a form of predator defence (Takeda,
2010) and can lower the chances of being preyed upon.
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Comparison of Individual and Group Behaviour | |
Similar to the individual
data, crabs groups of larger sizes took longer to bury and scurried further.
Scurrying distance was not found to be significant between individual crabs and
crabs in groups of 5. This suggests that when crabs swarm they do not act
differently than when crabs are found individually. However the burying time
between individual crabs and groups was found to significantly different
suggesting that crab confidence increases when surrounded by other individuals.
Supporting this, Murakami and colleagues (2014) showed that when soldier crabs
where in larger groups (15 crabs) they were more likely to cross a pool of
water then when in smaller groups (5 crabs). This suggests that soldier crabs
may find safety in numbers. Behaviour similar to this can be seen all over the
animal kingdom. For example fish are seen to shoal when predators are present (Pitcher,
1992), birds flocking in Hawaii (Hart
& Freed, 2005) and impala form herds in South Africa (Smith
& Cain, 2009).
There are many advantages
for congregating in groups such as group size discourages predators (Pitcher,
1992), lower predation on individual (Watt,
et al., 1997) and an increase number of eyes to identify predators (Smith
& Cain, 2009). However, discovered by Watt and team (1997) with a
study on tadpoles, the total strike right of predators on a group increased
with larger group sizes but the strike right per individual decreases. This
means individuals benefit but the group as a whole does not. Animals sometimes
have lower vigilance when in large groups (Smith
& Cain, 2009), though for the study this is not considered as M.
longicarpus burying time started once the crabs had identified the predator
and started to scurry away. As the scurrying distance did not vary amongst
individuals and groups, the soldier crabs stayed within the presence of a
predator for longer periods of time. Hence it is suggested that M.
longicarpus behave similar to the tadpoles when a predator is present. Thus
the individual is favoured and not the group.
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Limitations and Future Research | |
The greatest limitation
for the data was time. Only two days of field work were conducted and although
the study managed to find a significant difference for burying time between
individual and groups of crabs the standard deviation was large. Insignificant
variation were observed throughout all the data indicating that a larger sample
size should be to determine whether this is crab behaviour or is by random
chance. Acknowledging this, crabs behave different in various temperature and
climate conditions (Cameron, 1966; Kelemec,
1979), therefore to keep a fair test field work would have to be
conducted in similar conditions, as done for this study.
The group size of crabs
was only five as observers were unable to following each individual crab in
larger groups. Future research can be conducted using video cameras and image
processing software similar to the study by Murakami and team (2014). Using
image processing the scurry path and burying time of each individual would be
able to be recorded, thus larger groups of soldier crabs would be able to be
studied.
Other studies can
conducted to determine changing crab behaviour for individuals and groups such
as changing the sediment type and sand temperature.
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