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Prof. S. Ajmal Khan and Dr. S. Ravichandran
Centre of Advanced Study in Marine Biology
Annamalai University
rabs are among the most predominant species in many mangrove
C forests. Among the brachyurans, the grapsids, ocypodids and xanthids
are dominant. The sesarmids attain extreme diversity and richness in Indo‐
Pacific mangroves. They constitute about 80% of the macrofaunal biomass
and can reach densities of 80‐90/ m2. The crabs depend directly on
mangrove areas for survival, by feeding on leaves and litter. The crabs
have a significant role in detritus formation, nutrient recycling and
dynamics of the ecosystems, together with numerous annelids and
nematodes living in the sediment. The digging behaviour by crabs
enhances aeration and facilitates drainage of mangrove soils. They are
adapted to the sediment conditions, tidal fluctuations, and varying
salinities found in the mangroves. The crabs in the mangrove environment
show a variety of feeding modes. Two families of crabs are particularly
associated with mangrove ecosystem, the grapsidae and the ocypodidae.
Their ecological role is dependent up on the abundance of crab species in
the mangrove ecosystems. However, fewer publications have concerned
with the crabs in mangroves. Much more understanding on the taxonomy
and biodiversity of the group, which is also commercially important for
local human populations, is highly warranted for the threatened mangrove
ecosystems.
Crabs in the Mangrove Environment
Of all benthic macrofauna inhabiting the mangrove forests, grapsid crabs
are amongst the most important with regard to both the number of
different crab species and the total number of crabs present (Dahdouh‐
Guebas, et al. 1997). Most grapsid crabs belong to the subfamily Sesarminae
and many also belong to the largest genus in Sesarminae, called Sesarma.
Some of the more commonly mentioned species in the literature are
Neosarmatium meinerti, Neosamartium smithi, Sesarma messa, Sesarma
moluccensis, Metapograpsus latifrons, and Metapograpsus thukarhar (Smith
322 Brachyuran Crabs
1987, Lee 1998). There is considerable difference in the occurrence and
diversity of mangrove grapsid crab species depending on the particular
mangrove forest. The largest amount of diversity of grapsid crabs recorded
is from peninsular Malaysia where 51 species of grapsids, of which 44 were
sesarmines, were found (Lee 1998). This differs greatly from a recent review
of the grapsid crabs of the Americas where 23 species of Sesarma were
described but only 5 of these species are associated with mangroves.
Biodiversity study of crabs in the Pichavaram mangroves has shown that
there are 46 species of crabs from the five different stations (Ravichandran
and Kannupandi. 2007).
Brachyuran crabs, which contribute a bioenergetically significant
faunal component, play a significant role in maintaining a steady state of
the mangrove ecosystem. The diversity of the mangrove crabs is
exemplified by their wide feeding habits. The substratum, water level,
salinity, temperature and floral distribution were possible factors which
influence zonation and distribution of crabs in the mangrove environment.
The ghost crab Ocypode macrocera prefer sandy substratum and
Macropthalmus species prefer only muddy substratum. Uca species is found
in dry or grassy, elevated and muddy grounds. Grapsid crabs are the
dominant species among the crab taxa.
Grapsid crabs are considered significant seed predators of
mangroves and can be a threat to the successful regeneration or restoration
of mangroves. Although grapsid crabs tend to consume a large amount of
propagules, some studies have shown that they can be selective, selection
based on nutritive values of the propagules and the density of the specific
mangrove tree species in the canopy. Smith (1987) found a significant
difference in predation among the propagules of five mangrove tree species
which could almost entirely be accounted for by the difference in nutritive
values of the mangrove propagules. The leaves of Avicennia marina contain
rich nutrients and more palatable compared to other mangrove leaves (Lee,
1989; Rajendran and Kathiresan, 2000; Ravichandran and Kannupandi,
2004). This perhaps may be the reason for maximum number of grapsid
crabs present in the Avicennia zone.
In the mangrove environment, sesarmid crabs descend from their
burrows above the high tide mark to feed on mangrove leaves which are
then taken back into the burrow. Storage of mangrove litter within the crab
burrows however is influenced by ecological constraints rather than food
quality.
S. Ajmal Khan and S. Ravichandran 323
Importance of mangrove crabs
Crabs are good source of food to marine life as well as to man as a good
protein source (Khan, 1992 and Siddiqui et al., 2002). The nutritional quality
of the crab proteins compare very favorable than that of muscle meat of
mutton, chicken, duck and fish (Derosier, 1963; Newcombe, 1944; Zaitsev et
al., 1969). Besides, these crabs form the food for many birds, snakes and
predatory fishes. Brachyuran crabs are detritus feeders consisting of minute
particles of plant and animal origin. The detritus therefore is biogenic
material in various stages of microbial decomposition and represents a
potential energy source for consumer species. The faeces of the crabs
contain carbon, nitrogen, phosphorus and trace metals which form a rich
food for other consumers ( Kuraeter, 1976). Further, their burrowing habit
aids in aeration and free circulation of water which promotes the growth of
seedlings in mangroves. The burrowing members are of immense use in
recycling nutrients by “ploughing”. Grapsid and ocypodid crabs are
normally considered to be of no economic value but they assist in the
breakdown of particulate organic material by exposing them to microbes.
Their burrowing habit assists in oxidizing the sulphides that build up, due
to the high rates of organic decomposition in mangrove swamps (Diemont
and Van wijngarden, 1975). Degradation of mangrove leaf litter by searmid
crabs play a key role as a major link between primary and secondary
productions. Leaf consuming mangrove crabs play an important role in the
initial processing of litter in low to mid intertidal riverine and fringing
forests. Many sesarmid crabs are mainly herbivorous and play a vital role
in the degradation and thus in bio‐geochemical cycles. In this regard, keys
for identification are described here.
List of crab species reported in Pichavaram mangrove
S.No. Name of the species S.No. Name of the species
23. M. erato
(Source: Ravichandran and Kannupandi, 2007)
Methods of Collection and Preservation
Crabs are generally ‘easy to collect’ and most often hand picking is
very effective in intertidal and subtidal zones, while netting and trapping
are the commercial fishing practices. The burrowing intertidal crabs may
be collected either by digging or by pouring dilute formalin or weak acid
inside the burrow. In backwaters and estuaries, a smaller indigenously
made ‘ liftnet’ is used with a bait.
Crabs can be preserved wet in 6‐10% formalin neutralized with
hexamine i.e., 100 g per 1000 ml formalin. To avoid limb‐shedding the
unique phenomena of autotomy alive crabs may be norcotized first with
few menthol crystals or by adding few drops of chloroform and then
S. Ajmal Khan and S. Ravichandran 325
preserved in spirit of formalinated spirit for a day or two, to make the
autotomizing muscle as well their ‘breaking –planes’ rigid. After this fixing,
crab can be preserved in 6‐10% formalin for the laboratory studies.
Preservation of adult intermoult crab is ideal for taxonomical studies.
To prepare a dry exhibit for long term exposure the carapace and
the abdominal plate must be removed and glued back after scooping out
the flesh. Strong formalin should be injected through the joints of legs of
chelae. The carapace can be very easily removed with a simple technique.
For this crab could be held in left hand palm and yet your left thumb and
index fingers make a grip on the one side legs of the crabs. Then the
carapace can be pulled away by lifting the last anterolateral spine using the
right thumb and simply open the abdominal plate on the ventral side and
cut a portion with scissors to scoop out the flesh. Then the crab can be filled
with Borax‐an absorbent or dry sand and left for one or two hours under
any light source. When the crabs become free from moisture, the borax or
the sand can be brushed out. Strong formalin must be injected through the
segmental joints of chela and leg by a syringe. The carapace and abdominal
plates can now be glued back on its original position. The broken limbs or
the awkwardly shaped one can be broken and then wired by a thin copper
wire into position and glued with fevicol. Finally a coating with clear
varnish “sheenlac” makes the crab shinning.
The colour retension in crab is now experimented with 5% aqueous
solution of sodium arsenite to preserve red and related colours. Similarly
neutralised formalin with glycerine is felt better for the preservation of grey
or black coloured specimens. A mixture of 5 g sodium arsenite and 5 ml
formalin in 95ml water helps to retain white carapace colouration.
For taxonomical studies, the first pleopod or the third maxilliped
can be permanently mounted and the drawing made using the camera
lucida. Only the left pleopod or third maxilliped i.e. right as viewed from
ventral side of a crab can be removed. After treating with caustic potash
and staining in fast green, permanent balsam mounts can be made. While
mounting, it should be noted that if the gonopod is curved, the concave
border of the pleopod must be placed towards the outer side as a crab is
viewed ventrally. Similarly the distal three segments of the third maxilliped
namely propodite, dactylopodite and carpopodite must be facing inwardly.
Temporary mounts can be made with glycerine and alcohol. In addition to
camera lucida, a squared micrometer eye piece is also used to take
measurements.
326 Brachyuran Crabs
Designation of Brachyuran Crabs Regions
a. Dorsal View
1. Frontal 1. Merus of cheliped
2. Orbital 2. Carpus of cheliped
3. Progastrics 3. Propodus of cheliped
4. Mesogastrics 4. Dactylus of cheliped
5. Anterolateral 5. Merus of periopod
6. Metagastrics 6. Carpus of periopod
7. Epibranchial 7. Propodus of periopod
8. Mesobranchial 8. Dactyylus of periopod
9. Cardiac 9. Merus of natatory leg
10. Posterolateral 10. Carpus of natatory leg
11. Lateral post cardiacs 11. Propodus of natatory leg
12. Median post cardiacs 12. Dactylus of natatory leg
b. Ventral view of Male
c. Ventral view of Female
Ventral View
1. Suborbital
2. Pterygostomian
3. Third maxilliped
4. Antenna
5. Antennule
6. Sternum
7. Tergum
8. Telson
S. Ajmal Khan and S. Ravichandran 327
Key to Families of Mangrove Crabs
Carapace usually round or oval; right chela larger than left, last leg
flattened, adapted for swimming; usually a small lobe at inner angle of
endopodite of first maxilliped.
…………………...Portunidae
Legs not adapted for swimming; branchial regions not swollen; carapace
anteriorly broadened; no inner lobe on endopodite of first maxilliped
…………………. Xanthidae
Body squarish; carpus of third maxilliped not articulating with merus; last
legs not dorsally placed; front very narrow, orbits very elongate and
oblique covering almost anterior portion of carapace
……………….Ocypodidae
Last legs not dorsally placed; a gap between third maxillipeds; sides of
body straight or arched, front broad; rarely true land crabs
..…………………Grapsidae
Last pair of leg not dorsally placed; a gap between third maxilliped; sides
of body strongly arched, carapace transversely oval, front narrower; land
crabs
.…………Gecarcinidae
Key to Subfamilies of Family Portunidae
Four to nine anterolateral teeth; inner orbital distance broader; basal joint of
antenna usually broad and its anteroexternal angle lobulate; legs shorter
than cheliped, last pair typically paddle shaped
………………….. Portuninae
Two lateral teeth, inter orbital distance narrow; eye‐ stalk strikingly long
orbits occupying whole anterior border of carapace
………… Podophthalminae
Key to Genera of Subfamily Portuninae
A) Anteroexternal angle of basal antennal segment not appreciably
produced, flagellum thus standing in orbital hiatus
1. Anterolateral carapace cut into nine teeth of equal size;
propodus of chelipeds inflated
…………………………Scylla
328 Brachyuran Crabs
Posterior border of carapace straight and forming an angular junction with
posterolateral borders ……………... Goniohellenus
Sharp median lobule on lower border of orbit, no spine on posterior margin
of carpus of natatory legs ……………………Charybdis (Charybdis) lucifera
Posterior border of propodus of last leg serrated; spine on posterior margin
of carpus of natatory leg ………. Charybdis (Charybdis) helleri
Key to Species of Genus Thalamita
Spines of anterolateral border truncate, spines almost equal
…………… Thalamita crenata
Spines of anterolateral border square cut , last not enlarged
………..…. Thalamita chaptali
FAMILY : XANTHIDAE
Key to Subfamily, Genera, and Species of Family Xanthidae
Carapace usually much broader than long, transversely oval, some times
transversely hexagonal, front narrow, one‐ third to one fifth of the greatest
breadth of carapace
………………….. Xanthinae
Carapace granular marginally, regions vaguely define; basal antennal joint
not reaching front; anterolateral border with lobes or teeth
…............................... Galene
Carapace pentagonal, surface lumpy and scabrous near borders,
pterygostomian region almost hairy, anterolateral borders indistinctly four
lobed of which two distinct, posterolateral border longer than anterolateral
border; inner and outer angles of wrist spiniform
..……… Galene bispinosa
Carapace moderately broad; front about a third of the greatest breadth of
carapace; anterolateral borders of carapace not longer than posterolateral;
basal antennal joint does not or just touch front
…… Subfamily ‐ Piluminae
330 Brachyuran Crabs
Oblique ridges on palm high, predactile ridges parallel, ambulatory merus
narrow in both sexes
…..…Uca lactea annulipes
Merus of smaller cheliped posteriorly flattened and tuberculated; orbits
very oblique
… Uca triangularis bengali
SUBFAMILY : MACROPTHALMINAE
Key to Genus and Species of Subfamily Macropthalminae
Carapace broader than long, tooth at anterolateral angle of carapace
truncate and square cut …………….Macrophthalmus depressus
Carapace broader than long, front about a fourth of greater breadth of
carapace ……………..…….. …. M. erato
Key to Genus and Species of Subfamily Scopimerina
Carapace at least as long as broad; chelipeds nearly four times as long as
carapace; tympana present on sternum …. Dotilla myctiroides
FAMILY : GRAPSIDAE
Key to Subfamilies of Family Grapsidae
Front broad and deflexed; flagellum of antenna very short; external
maxilliped leaving rhomboidal gap …………. Grapsinae
Front broad and not deflexed, sublaminar; antennal flagellum lengthy;
external maxillipeds completely shut buccal cavern
.…………… Varuninae
Front broad and deflexed; external maxillipeds slender, transverse hairy
crest on ischium ..………… Sesarminae
Antennular fossets deeply divided into lobes; infra orbital border curved;
external maxilliped incompletely close buccal cavern
…………… Plagusinae
332 Brachyuran Crabs
Key to Genera and Species of Subfamily Grapsinae
Front less than half of extreme width of carapace; merus of external
maxillipeds longer than broad; finger of chelipeds spoon shaped at tip;
exognath of external maxillipeds well developed
……………….. Grapsus
Tooth at inner angle of wrist of cheliped straight
…....…Grapsus strigosus
Tooth at inner angle of wrist of cheliped curved
.…… Grapsus tenuicrustatus
Front more than half of extreme width of carapace; merus of external
maxillipeds broader than long ……………… Metapograpsus
Walking legs larger, dactylus distinctly shorter than propodus
.……………… M. maculatus
Walking legs shorter, dactylus nearly as long as propodus
………………….. M. messor
SUBFAMILY : VARUNINAE
Key to Genera and Species of Subfamily Varuninae
Exognath of external maxillipeds broader than or nearly as broad as
ischium; carapace very flat and lateral borders toothed, no postrolateral
facet defined
……………. Ptychognathus
Carapace minutely puncrate, two pairs of indistinct transeverse
depressions on carapace; upper orbital margins shaped
.…………….. P. altimanus
Exognath of external maxillipeds narrower than ischium; carapace
subcircular, teeth of anterolateral border flattish and separated by very
narrow notches
……………. Pseudograpsus
Legs hairy, not compressed, a fleshy lobe at base of fingers of chelipeds
…………… P. intermedius
S. Ajmal Khan and S. Ravichandran 333
SUBFAMILY : SESARMINAE
Antennal peduncle not excluded from orbit; posterolateral border of merus
of pereopod four to five with denticulation
..…………… Nanosesarma
Antennal peduncle not excluded from orbit; posterolateral border of merus
of pereopod four to five without denticulation , carapace with one or two
anterolateral teeth ……………. Neoepisesarma
Antennal peduncle not excluded from orbit; posterodistal border of
pereopod with out denticulation but no anterolateral tooth on carapace
……………… Parasesarma
Key to Subgenera and Species of Genus Nanosesarma
Outer palm of cheliped entirely covered with fur hair concealing lines of
granules …..……… N. (Nanosesarma)
Outer surface of palm of cheliped without pectinated crest; anterolateral
teeth on carapace clearly marked
……… N. (Nanosesarma) minutum
Outer surface of palm of cheliped smooth or with limited patch of hair,
merus of pereopod four with two to four acute spines posterodistally
……………… N. (Beanium)
Upper surface of palm of cheliped with two oblique pectinated crest, three
acute spines ( one long and two short) on posterodistal border of pereopod
four …… N. (Beanium) batavicum
Upper surface of palm of cheliped with numerous striae, one of which
forms pectinated crest, four strong spines on posterodistal border of merus
of pereopod four … N. (Beanium) andersoni
Key to Subgenera and Species of Genus Neoepisesarma
On upper surface of palm of cheliped low pectinated crest continued from
distal end to proximal end, numerous transverse long swollen dactylar
tubercles closely arranged in a continuous rim
…….. Neoepisesarma (Neoepisesarma)
334 Brachyuran Crabs
336 Brachyuran Crabs
Sesarma brockii Sesarma plicatum
Nanosesarma (Nanosesarma) minutum Grapsus strigosus
Neoepisesarma (Muradium) tetragonum Neoepisesarma (Neoepisesarma) mederi
Metapograpsus messor
Metapograpsus maculatus
S. Ajmal Khan and S. Ravichandran 337
Ptychognathus altimanus Cardisoma carnifex
Heteropanope indica Scylla serrata
Portunus (Portunus) pelagicus Charybdis (Charybdis) helleri
Portunus (Portunus) sanguinolentus Galene bispinsoa
338 Brachyuran Crabs
Ocypode macrocera Macrophthalmus depressus
Uca (Celuca) triangularis bengali Uca (Celuca) lactea annulipes
Sesarma (Beanium) andersoni Charybdis natator