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BIOTROPIA Vol. 16 No. 1, 2009: 1 - 10
*Corresponding author: jtmasagca@dasma.dlsu.edu.ph, jmasagca@yahoo.com.
FEEDING ECOLOGY OF TREE-CLIMBING MANGROVE
SESARMID CRABS FROM LUZON, PHILIPPINES
JIMMY TEVAR MASAGCA
De La Salle University-Dasmariñas
Cavite 4115, e Philippines
ABSTRACT
Despite the large ecological study of tree-climbing mangrove sesarmid crabs in other
countries, the Philippine representatives appear to have not been investigated extensively.
This paper presents the feeding ecology as to dependence on mangrove trees of sesarmids in
different mangrove areas of southern Luzon. This is biased on the nature of the crab habitats,
arboreal climbing skills and burrowing behavior of the sesarmids: Selatium elongatum and
Episesarma versicolor − exclusive mangrove tree climbers (EMTC); Sarmatium germaini −
occasional mangrove tree climber (OMTC); and the non-mangrove tree-climbing (NMTC)
sesarmids- Neosarmatium smithii, Perisesarma bidens and Perisesarma eumolpe.
Key words: mangrove crabs, sesarmid crabs, climbing skills, burrowing skills, arboreal climbing
crabs, Catanduanes, Philippines
INTRODUCTION
The crustaceans (e.g. brachyurans) are always a prominent and diverse element
of the fauna in mangals (Morton 1990). Crabs are diverse in mangrove habitats and
abundant in many mangals (Jones 1984). Several authors (e.g. Jones 1984, Lee 1998)
have attested that brachyurans are important in the mangrove ecosystem structure and
function. Unfortunately, it appears that there is still a dearth of detailed published
information on the occurrence of several families of this group of crustaceans in the
mangrove areas of the Philippines. Moreover, there seems to be limited published
data on the ways in which these faunal elements use the mangrove resources. The
sesarmids are less documented in the Philippines unlike in other countries, wherein
several reports are available from Singapore (Sivasothi 2000; Sivasothi et al. 1993;
Tan & Ng 1994; Ng & Liu 1999) Malaysia (Tan & Ng 1994; Leh & Sasekumar
1985), Indonesia (Soemodihardhjo & Soerianegara 1989; Rahayu & Davie 2002),
Hong Kong (Poovatchiranon 1986; Lee & Leung 1993; Lee 1998; Lee 1997; Kwok
1995; Kwok & Tang 2005; and Ashton 2002).
Crabs have different functions and impacts on the mangrove ecosystem. Firstly,
they can process as much as 70% of the leaf litter, and leaf processing can turn over
a litter at a rate in excess of 75 minutes (see studies of Leh & Sasekumar, 1985; Slim
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BIOTROPIA Vol. 16 No. 1, 2009
et al. 1997; Dahdouh-Guebas et al. 1999; Ashton 2002). Secondly, most mangrove
crabs feed on vascular plant materials, but they also feed on green leaves (Lee 1998;
Skov & Hartnoll 2002), including seedlings. Mangrove plant seedlings grazing,
particularly in the genus Sesarma may also slow regeneration of mangroves. On the
other hand, grapsoid crabs are the primary seed operators. Emmerson & McGwynne
(1992) described the feeding and assimilation of mangrove leaves by the crab Sesarma
meinerti. Moreover, the feeding ecology of Neosarmatium smithii was studied by
Giddins et al. (1986) and new records from Taiwan on this sesarmid crab was reported
by Naruse et al. (2006).
The crabs of the family Sesarmidae are known by various workers to have
significant ecological role in mangals (Lee 1998, Gillikin 2000; Gillikin & Schubart
2004, Gillikin et al. 2004). Many species of these mangrove crabs do not assimilate
much carbon from the mangrove leaves but rely on the sediments (Bouillon et al.
2002, Skov & Hartnoll 2002). Some species of the genus Perisesarma supplement
their diet with leaves (Leh & Sasekumar 1985, Slim et al. 1997). Hence, this aspect
on the feeding ecology of sesarmid crabs helps to trap the energy stored in these
leaves within the mangal before the tide can carry them away ( Lee 1998; Skov &
Hartnoll 2002).
Despite the large ecological study of mangrove sesarmids in other countries
(Bright & Hogue 1972; Hartnoll 1975; Micheli, Gherardi & Vannini 1991; Cannicci
et al. 1996; Slim et al. 1997; Dahdouh-Guebas et al. 1999; Fratini, Cannicci &
Vannini 2000, Flores et al. 2003) the Philippine representatives appear to have not
been investigated extensively. This study on the patterns of biodiversity in aquatic
systems of the Philippines attempted to determine the feeding ecology of selected
taxa of sesarmids as to (a) the nature of the crab habitats in the mangrove forest with
reliance to the mangrove trees, trunks, roots, and leaves that offer a wide variety of
ecological niches for the species under consideration; and (b) the behavior of the
sesarmid crabs as revealed by their burrowing abilities and tree-climbing skills needed
to explain the reliance of these crabs on the mangrove forest.
MATERIALS AND METHODS
Study sites
The collection and observation sites of the study include: I- Palsabangon
Mangrove Area in Pagbilao, Quezon, Philippines; II - Maqueda Channel in the Bicol
Region, Agojo Inlet Mangrove Reserve Area, San Andres and Palnab-Pajo Mangrove
Area, Catanduanes, Philippines.
Feeding ecology as to dependence on mangroves, arboreal climbing skills and
burrowing behavior
Specimens of mangrove crabs were collected by handpicking and with the use
of scoop nets and locally made traps. Collections were made both in the morning
and in the evening from the mangrove areas along rivers, creeks, inlets and the buffer
zones or marginal strips of the coastline in one site each for the study areas from June
2005 to February 2006.
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Ecological studies on feeding behavior took place in the purposively selected
study plots in Palnab-Pajo Mangrove Swamp, Catanduanes during the months of
October and December in 2005, while in the mangrove areas of Palsabangon in
Pagbilao, Quezon studies were done from January to February 2006. Ecological
studies of sesarmids were carried out in each of the mangrove areas following the
methods of Gillikin (2000). In the selected mangrove forests, presence or absence
of crab species were determined by visual inspection in 10 m diameter plots along
a transect perpendicular to the coastline, covering the full width of the forest. The
study investigated at least 10 plots along a 100 to 200 m long transect in the areas
under study.
Emergence and re-emergence of the sesarmids
During fieldwork, each plot was first inspected using binoculars from a distance
of approximately 6 m for 20 min. and the crab species recognized were noted and
recorded. Afterwards, naked eye observations were undertaken with two field workers
(observers) sitting on opposite sides of the plot for at least 30 min, following Hartnoll
et al. (2002) and Skov et al. (2002). As a modification, the researcher and field
assistants were positioned in two separate paddled boats (“banca”) when the plots
were observed. The researcher approached the plot and a disturbance was created
(either by throwing some mud, twigs or branches of mangrove plants) so that the
sesarmids, under study take refuge in the burrows, or holes, and other materials to
take cover (e.g. leaves, wood, twigs etc.). The time at which the first individual of
the sesarmids will emerge or re-emerge was recorded. Observations for this portion
of the study were undertaken during the day at low tides. Diurnal and lunar cycles
were not used in the distinction of the observation. The aforementioned procedures
are modifications of the works of Gillikin (2000), Hartnoll et al. (2002) and Skov
et al. (2002).
RESULTS AND DISCUSSIONS
Reliance on the mangroves as habitats
Table 1 shows the summary of the observations made on the habitats of various
grapsoid sesarmid crabs in the mangals of Catanduanes Island and Pagbilao, Quezon.
The nature of their habitat, climbing abilities and burrowing skills could explain the
feeding ecology of the mangrove crabs under investigation.
Table 1. Summary of the Feeding Ecology of Grapsoid Sesarmid Crabs from Different Mangals in
Luzon
Grapsoid Sesarmid
Species
Behavior/Nature of Habitat
Related to Feeding Remarks on fi eld observations
Episesarma versicolor Burrower and exclusive mangrove
tree climbing (EMTC) species
Migrating from burrows to the tree stems and
some at the canopies; constructs burrows around
soft sediments of mangrove roots and tree trunks;
seen feeding on calyx and leaves; seen cropping on
leaf litters and bringing fragments to the burrows.
Feeding ecology of tree climbing mangrove sesarmid crabs – Jimmy T. Masagca
4
Selatium germaini Occasional mangrove tree climber
(OMTC) species
Climbs on the branches of R. apiculata and R.
mucronata in the mangroves of Catanduanes.
Neosarmatium smithi Burrower and Non-mangrove
tree climbing (NMTC) species
Constructing burrows on mudfl ats;
Not seen climbing on stems of mangrove trees in
Quezon and Catanduanes
Perisesarma bidens Burrower and Non-mangrove
tree climbing (NMTC) species
Seen resting on Rhizophora tree (just within the
water lining) during high tide apparently for
avoidance and possibly for feeding purposes; also
seen hiding on rocks, crevices of boulders and tree
falls.
Perisesarma eumolpe Burrower and Non-mangrove
tree climbing (NMTC) species
On rare occasions, some crab samples were caught
on the trunks possibly carried by water current
during high tides.
Selatium elongatum Arboreal climber or EMTC in
mangrove trunks, branches and
canopies
Seen on main branches of Rhizophora; on aerial
roots; also on crevices of trunks.
Episesarma versicolor feed on calyx and leaves and crop on leaf litters, then
bring fragments to the burrows. The grapsoid sesarmid crabs, Perisesarma spp. and
Episesarma sp. were found to be herbivores and omnivore/deposit feeders, eating
mangrove litter and water plants. Moreover, the sesarmid crabs Sarmatium germaini
(Figure 1), Perisesarma eumolpe and Neosarmatium smithi feed on the mangrove litter,
composed of fallen mangrove leaves of Rhizophora, seedlings, calyx and twigs that
fall from the trees on the forest floor and into the water. This study confirmed that
the leaves of the mangroves, R. marina, R. apiculata, and Ceriops tagal were devoured
by the generally observed herbivorous/omnivorous crabs such as Perisesarma spp.
The feeding habits of mangrove crabs have been divided into seven groups by Jones
(1984): herbivore, carnivore, omnivore, deposit feeder, omnivore/deposit feeder,
specialized filterer, and filterer/omnivore. In the study of Islam & Uehara (2005),
stomach content analysis showed that P. bidens diet consists mainly of mangrove leaves
fragments, with small amounts of animal, algae and sediment matters, indicating that
this sesarmid is primarily detritivorous.
In Chiromanthes onychophorum, Malley (1998) noted that this sesarmid crab
consumes fallen leaves or their fragments, incompletely digests them, and returns
them to the environment as fecal matter in a more finely-divided state than when they
were ingested. Of recent, Ya et al. (2008) reported that both Perisesarma eumolpe and
P. indiarum are mainly sediment grazers, but also feed on mangrove leaves and roots
and occasionally animal matter. These crab species prefer Avicennia alba leaves to
other mangrove species, i.e., A. officinalis, A. rumphiana, R. apiculata and Bruguiera
gymnorhiza.
Dahdouh- Guebas (1997), Ashton (2002), Buck et al. (2003), Thongtham
& Kristensen (2003) and Schwamborn et al. (2006) analyzed the diets of sesarmid
crabs showed that their diet mainly consisted of mangrove leaves and in addition to
bustle animal matter.
BIOTROPIA Vol. 16 No. 1, 2009
Table 1. Continued
5
A. B.
Figure 1. Sarmatium germaini from Palsabangon Mangrove Area in Pagbilao, Quezon (A) and Palnab-
Pajo Mangrove Swamp, Maqueda Channel, in Bicol Region (Catanduanes) (B).
Burrowing and arboreal climbing skills of sesarmids
Similar with other sesarmids, P. bidens are burrowers, but observations made
in Quezon and Catanduanes indicate that some individuals tend to hide in natural
refuges found in the mangals investigated. These hiding places are the crevices of rocks
and boulders and in between portions of root buttresses of mangrove trees.
Field observation in Quezon, indicates that these crabs are also having preferences
in arboreal environments wherein Selatium elongatum and Episesarma versicolor were
seen clinging on mangrove stems or rush to the mangrove stems when disturbed,
while the boat is cruising in the waterways. These tree climbing crabs appear to be
less antagonized during the investigation and remained to be motionless for about 1
hour. When tides are rising or lowering, these crabs appear to be unaware as observed
for at least an hour. However, when the rising tide reaches their location in any of the
protruding branches of the mangrove trees in the embankments, these tree-climbing
crabs tend to adjust or move slowly with their heads upside down. Researches indicate
the possibility that these crabs cling to the mangrove stems not to find food but
as avoidance from very long immersion in the water during high tides and due to
the presence of predators. This could not be confirmed in the study. During field
observation, it was rather easy to find P. bidens in the open sand or mudflats.
Behavioral differences between the male and female samples of P. bidens was
observed. On the sexes, results of initial experiment suggest that there is no difference as
to the aggressiveness. However, the males are more receptive to antagonistic encounters,
especially when disturbed inside the aquarium prior to feeding observation. It was
difficult for the researcher to catch this sesarmid due to its swiftness and alertness.
One striking character of this crab is its fast action in retreating from external stimuli
such as touch or other mechanical disturbance. This could be the reason why the
feeding experiment did not proceed because the individuals of P. bidens placed in
the glass aquaria did not feed on the pieces of R. apiculata, R. mucronata, A. marina
and other associated mangrove plants.
However, the report of Ólafsson et al. (2002) cited that male and female crabs
are likely to have different energy requirements, due to differences in the energetic
costs of producing eggs or sperm (Michell 1993). A study of Kyomo (1992) conducted
on the sesarmid crab Sesarma intermedia showed that females were more specialized
in their feeding habits and had higher assimilation rates than males.
Feeding ecology of tree climbing mangrove sesarmid crabs – Jimmy T. Masagca
6
N. smithii form burrows around R. mucronata and R. apiculata roots but rarely
in Avicennia marina found in the mangals of Catanduanes. Observations indicate
that N. smithi feed on leaf litter but also some shrimps, specifically Palaemonetes sp.
This was observed during the afternoon hours when tides start to rise.
Observation on emergence and re-emergence of sesarmids in the burrows
In general, P. bidens appears to be the most active of the three sesarmids observed
during the day. As shown in Table 2, the red-clawed grapsoid sesarmid crab, P. bidens
emerged and re-emerged from the burrows much faster (mean of 3.48 mins) than
the sesarmids N. smithi (mean of 4.15 mins) and E. versicolor (3.93 mins). All three
sesarmid crab species display similar behavior of picking up the leaf fragments by
first standing still in motionless for about 7 to 10 seconds. Some sesarmids, e.g E.
versicolor has to consume the leaf fragment where they found it, then migrate for at
leas 0.5 meters and to be continued at the mouth of the burrows. Some crabs move
away from the areas where they got the leaf fragments and continue feeding near the
burrows. Some bring these fragments inside the burrows but mostly will have the
tendency to leave behind when disturbance was made.
There is much to be learned as to the sesarmid crab’s feeding ecology in the
Philippines since there are various similarities and differences in the findings of other
authors. Some researchers report on the unusual presence or occurrence as to their
level of dependence in the different habitats, arboreal or burrows. Climbing skills and
burrowing abilities also vary depending on the species. While others focus on how
sesarmids rush to the mangrove tree and and cling to the branches, it is still unknown
whether they are apparently resting or catching some insects or invertebrates. Other
researchers put premium on their occurrence in the mudflats or soft sediments of
mangals for feeding on some leaf litter. Moreover, the sesarmids under study have
been observed cropping and bustling on sand sediments and other fragments of
leaf litter that some crab ecologists might have ignored and do not give emphasis.
Further studies can be carried out in the future in terms of the feeding ecology of
these vital species of mangrove crabs in relation to morphological variation of the
feeding apparatus. Lastly, deeper reflections on the notions of convergent evolution
(Fratinni et al. 2005) and the timing of development in relation to arboreal climbing
skills and burrowing abilities of sesarmids deserve further attention.
Table 2. Data on the time of re-emergence of sesarmid crabs (P. bidens, N. smithi and E. versicolor) in
burrows after disturbance
Sesarmid Crab Species
Time of Re-emergence
MEAN TIME OF
RE-EMERGENCE (in
minutes)
TRIALS (Each consists of 3 repeats)
(in minutes)
123
Perisesarma bidens 3.35 3.65 3.45 3.4833
Neosarmatium smithi 3.95 4.05 4.45 4.1500
Episesarma versicolor 2.78 4.25 3.75 3.933
BIOTROPIA Vol. 16 No. 1, 2009
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CONCLUSIONS
Based on the results presented, Selatium elongatum and Episesarma versicolor are
exclusive mangrove tree climbers (EMTC); Sarmatium germaini as occasional mangrove
tree climber or OMTC; and the sesarmid species − Neosarmatium smithii, Perisesarma
bidens and Perisesarma eumolpe as non-mangrove tree climbers or NMTC. The sesarmid
species, E. versicolor, N. smithii, P. bidens and P. eumolpe posses burrowing skills, while
P. bidens appeared to be the most active of the three sesarmids investigated.
ACKNOWLEDGMENTS
The author acknowledges with thanks to the administration of De La Salle
University-Dasmariñas (DLSU −D) University Faculty Research Office (UFRO)
Director Dr. J. Morta; Education Dean, Dr. O. Legaspi ; OIC VVice Chancellor,
Dr. C. Cervillon; and DLSU−D Br. President Gus Boquer - for funding this research
project on the mangrove crabs. Sincere thanks is also given to Dr. David Gillikin, Vrije
Universiteit, T Brussel, Belgium and Dr. Peter K.L. Ng of the National University
of Singapore for sending reprints on the mangrove crabs and Ms. Marivene Manuel,
Philippine National Museum (PNM) for evaluating the earlier drafts of the report on
this study. My special thanks to Elver Sison, while finalizing this paper. The assistance
of my brother (Rico Masagca ); sisters (Tersy, Rosie, Elsie, Margie), and aunt (Mrs.
Florida Tevar) is greatly appreciated during field observations in Catanduanes. Special
thanks to the anonymous referees.
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