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FOOD AND FEEDING ECOLOGY OF FIDDLER CRABS SPECIES FOUND ALONG

THE COAST OF PAKISTAN

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 FOOD AND FEEDING ECOLOGY OF FIDDLER CRABS
SPECIES FOUND ALONG THE COAST OF PAKISTAN

NOOR US SAHER* , NAUREEN AZIZ QURESHI**

The food and feeding behaviour of four fiddler crab species commonly inhabiting
Sandspit and Korangi creek mangrove areas in Pakistan were investigated (50 specimens per
species). The mean number of spoon tipped setae on the second maxillipeds ranged
from 111 to 247 and were the highest in Uca iranica followed by U. annulipes, U. urvillei
and U. sindensis. The number of spoon tipped setae on the second pair of maxillepeds
did not differ between the sexes, but did differ among species (F3, 199 = 31.9, P < 0.005
and F3, 199 = 25.8, P < 0.005, respectively). The foraging distance away from the
burrow, and the shape of the radiating paths were species specific and positively
correlated with the diameters of burrows. The relative percent mud content in the
stomach was significantly different among the four species. Stomach contents of all
four species largely consisted of the plant debris, different interstitial organisms, their
eggs, spines, animal tissues, and filamentous algae, including organic detritus. U. urvillei is
the only species that was observed as predator species.
Keywords: Fiddler crab, feeding path, food, Spoon tipped setae, Pakistan.

INTRODUCTION

Foraging activities of fiddler crabs produce measurable changes in the

sediment structure and characteristics (Posey, 1987; Botto & Iribarne, 2000). These
crabs along with members of grapsids are considered as ecosystem engineers as
they continuously modify the surface sediment structure through their burrowing
and feeding activities (Teal, 1958; Kostka et al., 2002). Fiddler crabs are surface
deposit-feeders and forage upon the substrate (Macintosh, 1984; Robertson &
Newell, 1982) and are important consumers of detritus, bacteria, fungi, and benthic
micro algae in coastal marsh, mangrove, sand flat, and mudflat habitats (Backwell
et al., 2006; Mokhlesi et al., 2011). Their main food resource is organic matter,
including the endofauna sorted out from the substrate (Murai et al., 1982). The
unwanted sediment material left from sorting is formed into irregular cohesive
masses called food pellets or mud balls that are deposited on the surface sediment
near the burrow openings.
In decapods, the feeding ecology in their natural environment is frequently
determined directly from an analysis of their stomach contents, and the structure
and function of the mouthparts (Hill, 1976; Williams, 1981; Choy, 1986; Chande &
Mgaya, 2004; Hegele Drywa & Normant, 2009). The frequency of feeding and the
quality of the food ingested depend on the morphology of mouth parts, locality

ROM. J. BIOL. – ZOOL., VOLUME 59, No. 1, P. 35–46, BUCHAREST, 2014

36 Noor Us Saher, Naureen Aziz Qureshi 2

inhabited (Takahashi & Kawaguchi, 2001; Turra & Denadai, 2003). Actual food
supply depends on the productivity of the ecosystem, microbial activity, substrate
texture, and tidal action (Twilley et al., 1995; Moura et al., 2000) such that the
optimal conditions for foraging can be affected by texture, organic matter and
water content of the sediment (Reinsel & Rittschof, 1995). A comparison of local
population in different feeding environments reveals correlations among food
habits, feeding behaviour and feeding organs, which reflect the adaptation of
organisms to the food sources, the process of adaptation, and possibly speciation in
fiddler crabs (Colpo & Negrieros-Fronsozo, 2003).
Female crabs possess two isomorphic feeding chelipeds, and male fiddler
crabs are equipped with dimorphic chelipeds comprising of one small chelae
mainly use for feeding and one enlarged hypertrophied or major cheliped. The
feeding process involves two organs, the small chelipeds (minor cheliped in case of
male) that are used to scoop up sediments from the surface of the substrate and
transfer these to the mouth or buccal cavity, and the other are the spoon tipped
setae of the second maxillipeds that perform the function of sorting of food
material for ingestion (Miller, 1961; Ono, 1965; Crane, 1975; Robertson & Newell,
1982 a, b). The food particles are sorted by water passing through the buccal cavity
that suspends fine particles, which are trapped among the setae on the feeding
appendages (Ono, 1965; Crane, 1975; Robertson & Newell, 1982a). The ‘spoon
tipped’ setae have broad tips that are cupped and arched inwards (Lim, 2004). The
morphology of the minor cheliped is also adapted to the substrate texture; its frame
shape differs between fiddler crabs inhabiting mud and sand (Crane, 1975; Icely &
Jones, 1978; Rosenberg, 2002).
The objective of this study was to investigate the food and feeding behaviour
of four species of fiddler crabs found along the coastal line. There is not much
information on the feeding ecology of fiddler crabs commonly inhabiting the mangrove
areas of the Pakistan Coast. This is the first attempt to investigate the feeding
biology and ecology of fiddler crabs along the coast of Pakistan. Present study
mainly based on the hypothesis that the food preference is found among the species
and vary among the four species of fiddler crabs as preferred habitats of these
species varying in tidal height, vegetation and sediment composition.

MATERIAL AND METHODS

Study site
Two sites, the Sandspit and Korangi creek, were selected for studying the
food and feeding behaviour of the fiddler crabs. The Sandspit back waters
mangrove area (24o50’N, 66o56’E) is located in the south west of Karachi and is
connected to the Arabian Sea through the Manora Channel. The Sandspit beach is
bifurcated by a dry strip of land, with mud flats and dense stands of the mangrove
Avicennia marina on the northern side, and of sandy beach on the south side. The
3 Food and feeding ecology of four Uca species 37

Korangi creek mangrove area (24o79’N, 67o20’E) near the salts works in the
fishing village, Ibrahim Hyedri. Korangi Creek is the northern most creek of the
Indus Delta and connected at its northeastern end with Phitti creek and Kadiro
creek, while at its southwestern end, it connects with open sea and with Gizri
creek, and is bounded on its sides by extensive mangrove vegetation of A. marina.

Methodology
The shapes of feeding pellets and their distribution pattern around the burrow
were observed for three species of fiddler crab (Uca annulipes, U. sindensis and U.
iranica). The foraging distances (Fig. 1) of at least six pathways were measured
and correlated with the burrow diameter (BD). It was not possible to study the
feeding ecology of a fourth study candidate, U. urvillei, as this species is usually
found under the canopy of mangrove vegetation in moist swampy soil that
collapse, making the evaluation of foraging track and the shapes of feeding pellets
unreliable. Male U. urvillei at the study sites can wander many meters from their
burrows and sometimes predate on small shells and crabs.

Fig. 1. Foraging tracks of three species of fiddler crabs

(Uca iranica, U. annulipes, U. sindensis) at the study sites.
38 Noor Us Saher, Naureen Aziz Qureshi 4

It was observed that the male Uca crabs usually use their one small chelipede
for feeding and female use the both pincers. It was also observed does the use of
one sided chelae for feeding in male crabs effect the directional use of male crabs
mouth parts for the processing of food or the handedness makes any correlation
with the feeding process and organs involved in the process. In the laboratory, the
left and right second maxillipeds of each crab were placed on a cavity slide. The
presence and numbers of spoon tipped setae were counted under the compound
microscope and the differences in numbers of spoon tipped setae between the left
and right side of the maxillipeds were compared. Two factorial analysis of variance
(ANOVA) tested if the numbers of spoon tipped setae varied with sex or among the
species.
For the stomach content analysis about 50 specimens of each species of four
fiddler crab species (U. annulipes, U. sindensis, U. iranica and U. urvillei) were
randomly collected during the low tide period. The collected crabs (n = 200) were
immediately fixed in 10 percent formalin in the field and transferred into 70%
alcohol after 24 hours. Each crab stomach contents were examined.
Firstly a visual estimation of the percent gut fullness based on an index of 1–4,
with 1 equal to 0 to 25% gut fullness and 4 equal to 75 to 100% fullness of the gut
was made before opening dissecting out the contents into a petri dish. The percent
occurrence of mud or sand was categorized 1: 10%, 25%, 50%, 75% and 90%. The
stomach contents were washed with alcohol in a petri dish and the food categories
(eggs of invertebrate, crustacean appendages, nematode worms, plant tissue debris,
spines, animal tissues, and filamentous algae, etc.) were identified under a compound
microscope. The occurrence of the food categories quantified using the percentage
occurrence method (Williams, 1981; 1982), frequency of food item/number of
crabs’ %. This method gives a measure for the regularity with which food has been
taken in the highest value as the most important one in the sample or population.

RESULTS

Lengths and shapes of foraging paths

One way analysis of variance (ANOVA) showed a significant difference
between the three species of fiddler crab in the lengths of foraging tracks (F 2, 363 =
13.61, P < 0.001).
In U. sindensis the shape of the feeding pellets was biconvex and rounded
(Fig. 1). The mean feeding passage distance was 77 ± 25.4 mm with an average
burrow diameter of 10.2 ± 2.9 mm. Positive linear relationship (r2 = 0.82) was
observed between the size of burrow diameters and an average distance of feeding
passages around the burrows.
5 Food and feeding ecology of four Uca species 39

The feeding pellets of U. annulipes were usually crudely rounded in shape.

The mean feeding passage distance was 99 ± 36 mm with the mean BD of
11 ± 2.6 mm, the relationship between the BD and the feeding passage distance
was also positive (r2 = 0.6).
The feeding pellets of U. iranica were barrel in shape. The mean feeding
passage distance of U. iranica was 102 ± 37.8 mm with an average BD of
10.2 ± 3.2 mm (Table positive relationship (r2 = 0.66) was observed between BD
and the average feeding passage distance around the burrows.

Spoon tipped setae

The number of spoon tipped setae separately showed a significant difference
among species, but they were not significantly different between the sexes (F3,199 =
31.9, P < 0.005 and F3,199 = 25.8, P < 0.005 respectively). Post hoc analysis showed
that the spoon tipped setae were significantly highest in U. iranica, compared to
U. annulipes which were in turn significantly greater than U. urvillei and U. sindensis
(U. ira. > U. ann. > U. ur. > U. sin.).

Gut content analyses

Initially the gut fullness of the specimens of each species collected for
analysis was observed through binocular visual estimation (Fig. 2). Gut contents of
all four species were largely comprised of plant debris, different interstitial organisms
and their eggs and organic detritus (Fig. 3). The percent proportions of mud in the
stomachs was significantly different among the four species (F3,199 = 18.29,
P < 0.005). The highest mud content was found in U. urvillei followed by
U. sindensis (Fig. 3).

Fig. 2. Percent gut fullness estimated in the four species of Uca.

40 Noor Us Saher, Naureen Aziz Qureshi 6

U. urvillei
40 U. annulipes
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Fig. 3. Percent frequency occurrence of the various food items identified

in the stomach of four Fiddler crab species.

The diet of U. annulipes was mainly based on the plant tissue debris and
filamentous algae and the leaves of Avicennia marina (Forsk), identified on the
basis of tissue and stomata feature. U. sindensis and U. iranica had invertebrate
and crustaceans and animal remains in their diets. U. urvillei rarely fed on fresh
leaves of mangrove trees and displayed opportunistic diet, the male crabs were
observed wandering many meters from their burrows, predating small crabs and
molluscs by crushing their shells with the major cheliped; they were also seen to
feed on surface algae. This is reflected in the significant difference in the ratio of
animal to plant matter in the stomach contents of each species.

DISCUSSION

The three species of Uca (U.iranica, U. sindensis and U. annulipes) were

observed to forage around their burrows in circular or radiating paths. Crabs that
forage moving in a circular area, conserve the time and distance travelled away
from a burrow, for a given total area harvested. Radiating circular paths optimize
7 Food and feeding ecology of four Uca species 41

net resource yields, especially when resources are homogeneously distributed

(Smith, 1968; Covich, 1976; Andersson, 1978; Hixon, 1980, Schoener, 1983). It was
also observed that crabs maintain their positions by orienting to the pellets and
subsequent troughs that are created as they forage. The arrangement of feeding
pellets around the crabs burrow showed interspecific variability. Mukhlesi et al.
(2011) observed the intersexual difference in the U. sindensis during their mesocosm
studies. Circular paths erase the selection among food concentrations, especially
when the burrow provides an essential or limiting resource constraining foragers to
balance conflicting demands (Zimmer-Faust, 1987). The burrows provide crabs
with their only reliable source of water during an ebb tide. A resource that is
essential to continue feeding activity and to resist desiccation. Systematic grazing
also reduces the chance to feed in previously grazed areas and it was commonly
observed among other invertebrate and vertebrate grazers (Davies & Houston,
1981; Waddington & Heinrich, 1981). The shapes of the feeding pellets also varied
in three species studied, U. sindensis, U. iranica and U. annulipes. This difference
can be related to the shape and the depth of the spoon tipped setae, arrangement of
spoon tipped setae on maxillepede, and structure of small cheliped and these
observations indicated the need of a detailed study.
Spoon tipped setae on the second maxilliped decreased in abundance in the
following order: U. iranica > U. annulipes > U. urvillei > U. sindensis. Uca iranica
and U. annulipes were found in sandy habitat and U. urvillei and U. sindensis
inhabited muddy areas. Ocypodidae inhabiting sandy habitat generally have more
numerous spoon tipped setae on the inner surface of the second maxilliped and the
outer surface of the first maxilliped compared to those inhabiting muddy flats,
which have a few spoon tipped and many plumose setae (Miller, 1961; Ono, 1965;
Icely & Jones, 1978). Lim (2004) compared spoon tipped setae in two species
U. annulipes and U. vocans and also found more spoon tipped setae in U. annulipes
which inhabits the sandy habitat.
The number of spoon tipped setae differed between sexes. Weissburg (1991)
found quantitative differences in spoon tipped setae of male and female U. pugnax.
Lim (2004) found a difference in the area of maxilliped that was covered by spoon
tipped setae and a number of setae and observed no significant difference of spoon
tipped setae between male and female crabs of U. vocans and U. annulipes.
The food composition of Decapoda in their natural environment is usually
determined directly from an analysis of their stomach contents, even though
identification of the food remains is difficult. This is due to the structure and
function of the mouthparts (Hill, 1976; Williams, 1981; Chande & Mgaya, 2004).
In the present study, the stomach contents analyses of four species of Uca
revealed that they primarily feed on the plant debris, different interstitial organisms
and organic detritus. The eggs of invertebrates, crustacean appendages, nematode
worms, plant tissue debris, spines, animal tissues, and filamentous algae were the
main food categories identified from the gut material of four species of Uca. These
results show that the species of fiddler crabs were identified as omnivorous, feeding on
detritus, animal and plant matter. The three studied species (U. annulipes and
42 Noor Us Saher, Naureen Aziz Qureshi 8

U. urvillei) were reported as algal feeders (Icely & Jones, 1978) but in the present
study the presence of various food items including algae indicates the studied Uca
species can be considered as omnivorous rather than algal feeders. Omnivorous
organisms with more feeding options have greater ability to optimize nutritional
balance compared to ones with narrower diet breadth (Bjorndal, 1991).
Table 1
Mean, minimum and maximum number of spoon tipped setae on the left and right maxillipeds
of four species of fiddler crab (STS = Spoon Tipped Setae)
Variable Crab species N Mean± SD Min Max
STS ON LEFT
MAXILLEPEDE
Uca urvillei 5 157.8±26.3 12 196
Uca sindensis 5 102.3±11.9 85 119
Uca annulipes 5 210.5±65.4 82 302
Uca iranica 5 247.1±48.8 147 320
STS ON RIGHT
MAXILLEPEDE
Uca urvillei 5 162.4±30.7 110 210
Uca sindensis 5 111.5±18.6 84 137
Uca annulipes 5 207.3±64.7 74 298
Uca iranica 5 246.8±41.8 156 310

Table 2
Summary of descriptive statistics for the foraging distance around their burrow
and burrow diameter of three species of fiddler crab
Variable Species N Mean± SD Min. (cm) Max. (cm)
Feeding
distance (cm)
U. sindensis 252 7.7 ± 2.5 4.0 14.0
U. iranica 279 10.2 ±3.7 4.5 17.5
U. annulipes 243 9.9 ± 3.6 4.5 19.0
Burrow
diameter (cm)
U. sindensis 50 1.02 ± 0.3 0.5 1.5
U. iranica 50 1.02 ± 0.3 0.4 1.6
U. annulipes 50 1.11 ± 0.4 0.6 1.8

Table 3
The presence of food content in the stomach of Uca species along the coast of Pakistan
(+ very few, ++few, +++ Numerous)
Food content Uca urvillei Uca sindensis Uca iranica Uca annulipes
Percent mud 50%–90% 40%–75% 5%–50% 10%–60%
Filamentous algae ++ +++ + ++
Plant debris +++ + +++ ++
Eggs – ++ +++ +++
Crustacean appendages + + + +
Spines +++ + ++ ++
Nematodes worms ++ +++ +++ +
Muscles ++ + + +
Other material + + + ++
9 Food and feeding ecology of four Uca species 43

According to Marshall & Orr (1960), most crustaceans are omnivorous,

including fiddler crabs ingesting the available organic food particles available on
the moist marsh surface. Shanholtzer (1973) analyzed the stomach content of
U. pugnax and revealed that one third diatoms, one fourth fungi, one fifth unknown,
one fourteenth vascular plants and small amounts of arthropod parts and for-
aminifers were found in the gut. The diet of Uca species studied may contain greater
amounts of the detritus-microbe complex. The detritus-microbe complex is widely
believed to provide a suitable energy source for detritivores. A variety of micro
flora and micro fauna living on these detrital particles are suitable food for fiddler
crabs (Darnell, 1967). Uca may supplement their protein intake by ingesting bit of
carrion, arthropods and nematodes (Darnell, 1967; Montague, 1980). This highly
opportunistic feeding habit, known also for West Indies Grapsidae (Von Hagen,
1977), seems to be a common feature of mangrove crabs. The presence of plant
material in the diets likely indicates that multiple type of food provides complementary
nutritional resources for fiddler crabs. It has been reported that crustaceans
frequently select small and medium-sized high-energy food items from which the
nutrients were readily assimilated (Morales & Antezana, 1983; Juanes, 1992;
Kennish & Williams, 1997).
Mud content in stomachs was highest in U. urvillei. Shwartz & Safir (1915)
believed fiddler crabs to have a feeding preference for mud high in algal biomass.
Skov & Hartnoll (2002) suggested sesarmid crabs might feed on mangrove mud to
supply their diets with microbial nitrogen on decaying leaf litter fragments. It has
been suggested that stomach contents often reflect the availability of food in the
environment rather than an animal’s Preferences (Hegele Drywa & Normant,
2009). Large male crabs of U. urvillei were observed predating small gastropods by
crushing their shells, similar observations have been made for the omnivorous crab
Hemigrapsus sanguineus which prefers small bivalve prey as well as for Armases
cinereum (Buck et al., 2003). The choice of food to be consumed depends not only
on its availability in the environment, but also on its assimilability sensu lato. It has
been reported that crustaceans frequently select small and medium-sized high-
energy food items from which the nutrients are readily assimilated (Juanes, 1992;
Kennish & Williams, 1997).

Acknowledgments. This study was funded by the Pakistan Science Foundation project PSF/Res/S3
KU/Envr (51) grant to NAQ and is gratefully acknowledged.

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*
Received September 10, 2014 Centre of Excellence in Marine Biology,
University of Karachi,
Karachi 75270, Pakistan
noorusaher@yahoo.com
**
Department of Zoology
Govt. College, University Faisalabad, Pakistan

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