Biogeographic Distribution of Hermit Crabs in San

Agustin West, Isla Verde, Batangas City, Batangas,

Philippines
Mempin, Czarina Mae R.1, Shara Mariz S. Santana1
and Ruthela P. Payawal1
1
Department of Biology, College of Science, Polytechnic University of the Philippines, Philippines

Abstract
Biogeographic studies practice a share on the conservation management and provide relevance in understanding several ecological
impacts acting upon populations. Hermit crabs successfully inhabited intertidal area to terrestrial niches. Researchers were able to conduct a
biogeographic study about these hermit crabs in the Verde Island Batangas. Specimens were collected through hand pick and were preserved
in a 70% ethanol and were identified through its morphology and color pattern. There were a total of 594 hermit crabs collect ed in Verde
Island in both sampling time comprising of 3 families namedly: Coenibitidae, Diogenidae, and Paguridae. The family Diogenidae contained
more species but family Coenobitidae had more individuals. Coenobita rugosus H. Milne Edwards, 1837 was the most abundant species, with
285 specimens, followed by Clibanarius virescens Krauss, 1843 with 132 specimens, Coenobita cavipes Stimpson, 1858 with 88 specimens.
Presences of these intertidal hermit crabs rely on its geographic distribution in the Philippines. High abundancy of intertid al hermit crabs
(Diogenidae & Paguridae) was recorded in Sitio Mahabang Buhangin than in Sitio Sampalocan. These finding show that Verde Island is an
important area for establishment and maintenance of several paguroidean species. Despite of being constantly trawled by local fishermen and
became hotspot as a tourist attraction, this area can still conserve its potential in terms of paguroidean abundance and thus, marine
conservation efforts must be given attention to maintain paguroidean species richness. Changes in temperature vary the abundance of intertidal
hermit crabs based on their behaviours; pH, however, researchers can’t tell whether there are changes in their behavior in rising and lowing
level of pH since the range of collected pH values is small.

Keywords: Biogeography, Paguroidean, Coenibitidae, Diogenidae, and Paguridae

Introduction activities such as fishing, tourism and navigation,

Biogeography is the study of how
organisms are dispersed over the Earth (Brown &
Gibson, 1983). The processes within studying
biogeography distribution and evolution on remote
islands includes large scales of time and space,
and this study remains to be the most difficult to
study and understand (Carpenter, 2005). It was
derive from two very general features of nature, as
we move along any geographical dimension,
environmental conditions tend to vary in a
predictable manner individual, species differ in
their abilities to adapt in this changes which
influence three biological processes, immigration,
extinction and evolution (Lomolino, 2000). Verde
Island located in Batangas City, Batangas,
Philippines. Indo-Malay-Philippines Archipelago
which has been long considered as the world’s
highest marine biodiversity. These islands are lies
within SSME or known as Sulu-Sulawesi Marine
Ecoregion situatued at the apex of the Coral
Triangle, or the world’s center of marine
biodiversity. Within this are lies the Verde Island
Passage, identified by Carpenter (2005) as one of
the rich biodiversity areas in the country. Verde
Island as Center of the Center of Marine
Biodiversity faces serious threats from human
land use, pollution. As for this, different
Philippines laws were developed to protect and
maintain the area’s biodiversity.
Intertidal or the littoral zone are thought to
be the most dramatic ecotone of all as what Reese
(1969) said. Organism lived in this area must
overcome different biotic factors that are affiliated
in this area such as temperature fluctuation,
dessication and exposure to salt and fresh water.
Hermit crabs have a key role in the marine trophic
chain (Fransozo et al. 1998). Hermit crabs are
known to be as the most successful marine
invertebrate for they have inhabited sublittoral up
to terrestrial zone (Greenaway, 2003). They serve
as predators, scavengers, detritivores and even
filter-feeders. Presence of these macroinvertebrate
animals and their interaction can enhance the
biodiversity of their habitats in the present and
most probably in the past (Greenaway, 2003;
Schembri, 1982, & Reiss et al., 2003). However,
there is a scarcity in knowledge about its
biogeography and taxonomy most notably in some
islands in the Philippines especially in Verde
Island. This study seeks to find out: (1) what are
the hermit crabs that can be found in Verde
Island? (2) Their frequency and abundance; and
1
(3) the factors that determines their presence.
Materials and Methods
Study area and sampling
The study area includes the coastal areas
of Verde Island, Batangas City, Batangas,
Phillipines. Two intertidal areas were sampled:
Sitio Sampalocan and Sitio Mahabang Buhangin
in San Agustin West. In each intertidal area, 3
substrates were chose based to their physical
feature: rocky substrate, characterized mainly of
solid rocks and offers a rough environment for
organism to live in; sandy substrate, soft shores
that consists of loose deposits of sand, empty
shells, and coral rubbles formed by deposition of
particles that have been carried by water currents;
and seagrass substrate where various seagrasses
grow. Seagrasses are
flowering plants that dwell in shallow salty and
brackish water. They are sensitive to changes in
water quality which makes them a significant in
indicating the overall condition of the coastal
ecosystems.
The sample period was during the months
of November (Wet Season) and January (Dry
Figure 1. Map of Verde Island, Batangas City, Batangas, Philippines (Google Map, 2018)
Season). Hermit crabs were collected through
hand picking covering 60m latitudinal and 10m
longtidunal in both Sampalocan and Mahabang
Buhangin in each substrate. The hermit crabs were
removed from the shells. Total length (TL) and the
cephalothoracic shield (SL) were measured with a
ruler (10 mm) and was preserved in a 70% Ethanol
(Moradmand, 2007).
Construction of Dichotomous Key
Construction of dichotomous key was
made in order to identify the collected specimens.
It was made based on morphological and
distinguishing characteristics each specimen
possesses and contains concepts such as shield
length (SL), appendages, pereopods, and scales.
Included in it were figures such as photographs
and the defining characteristics. Photographs and
initial classifications were emailed to other
institution for the validity of nomenclature.
Physico-chemical Analysis
In order to correlate the collected intertidal
hermit crabs with the different environmental
factors;
2
physico-chemical analysis was conducted. Physico-
chemical parameters including pH and temperature
of sea water were measured using pH meter and
water thermometer respectively.
Species Distribution
In the field of ecology, the relative
abundance distribution (RAD) or species
abundance distribution describes the relationship
between the numbers of species observed in the
field study as a function of their observed
abundance. Correlations of environmental factors
were calculated using the software Paleontological
Statistics ver. 1.88.
Results
Taxonomic Identification and Geographic
Distribution
Family Coenobitidae
Coenobita brevimanus Dana, 1852
The material examined for C. brevimanus
specimen has a total length (TL) 163 mm and
shield length (SL) up to 100 mmIts ocular
peduncles are sub-cylindrical with terminal cornea
and the antennal peduncle not merged with the
second segment. Specimen displays purple
coloration, faint on thorax area, yellowish color on
dactylus of 2nd and 3rdpereopods. Chelipeds of C.
brevimanus are dissimilar and unequal, larger and
broader on the left while pereopods 1-3 have small
spines and have dark tips except for lighter tipped left
chela. In addition, its soft abdomen are spirally coiled
and it is covered with a number of setae on dactylus of
pereopods, partially obsolete on outer lateral side of
merus. Right antennal flagellum is lighter in color than
the rest of its dark-colored antennal flagellum.
Antennular peduncle and flagellum have orange and
brown coloration. A distinctive characteristic of C.
brevimanus is the presence of brush of setae on the
inner surface of right chela only.
Coenobita rugosus H. Milne Edwards, 1837
Coenobita rugosus is closely similar with C.
cavipes. Both demonstrates dissimilar and unequal
chelipeds, left being the biggest, truncated tips of the
antennular flagellum, and dent separated by sharp ridge
on inner surface of prodopus of left 3rdpereopod, C.
rugosus can be separated with C. cavipes by the
presence of stridulating apparatus on the outer surface
of left chela, distinctive for C. rugosus species. C.
rugosus only has setal brush on upper part of chela
while C. cavipes possess setal brush on both. Pereopod
1-3 have black tips apart from for the left cheliped.
Material examined has a total length (TL) of 45 mm
and shield length (SL) up to 23 mm.
Coenibita cavipes Stimpson, 1858
Coenobita cavipes is a part of Coenobitidae
family wherein crabs live in terrestrial or semi
terrestrial environment. They are known for brown
coloration on pereopods and brown markings on th
3
 Table 2. Summary of Bigeographic Distribution of Intertidal Hermit Crabs (Diogenidae & Paguridae)

Taxa Cebu Bohol Balicasag Palawan Mindoro Panglao Pamilican

Family Diogenidae
Calcinus gaimardii P P P
Calcinus latens P P P
Clibanarius cruentatus P P P
Clibanarius ransoni
Clibanarius virescens P P P
Dardanus deformis P P P P P P
Dardanus lagopodes P P P P P
Dardanus megistos P P P P P
Family Paguridae
Pagurus aff moluccensis P

4
shield. Distinct for the presence of setal brush
on both outer surface of chelae. C. cavipes’
chelipeds are distinctly dissimilar and unequal,
left cheliped larger than right. One of
Coenobitidae family’s distinguishing
characteristic is the truncated tips of the
antennular flagellum. Material used in this
species measures 45 mm (TL) and its shield
length (SL) 25 mm. It has a short antennal
flagellum with spirally coiled soft abdomen and
ocular peduncles that is strongly compressed
laterally. Pereopods 1-3 are black tipped.
Family Diogenidae
Genus Calcinus, Dana 1851
Calcinus gaimardii (H. Milne Edwards, 1848)
Calcinus gaimardii has an orangey to
dark reddish background with purple coloration
on pereopods; terminal cornea with reddish and
short orange ocular peduncle, left eyestalk a
little longer than right. The right cheliped has 1
spine distally on lower outer margin of merus.
The 2nd and 3rd pereopods are coloured but
without bands. The Material Examined has a
total length (TL) of 35 mm and shield length
(SL) of 20mm. Calcinus gaimardii resembles
Calcinus morgani. C. gaimardii has an orange
coloration on ocular peduncle while in C.
morgani, it has a distal blue band and broader
ocular peduncle. Observed specimen has a
strong orange to brown shield, which is a
characteristic of C. gaimardii while the while
the entirety of the shield of C. mogani is light
colored.
Calcinus latens (Randall, 1840)
The observed specimen has a total
length (TL) of 40 mm and shield length (SL) up
to 20 mm. Calcinus latens can be recognized
having an orange colored band on its prodopus.
One of characteristics of Calcinus species is that
the left eyestalk is longer than right. Chelipeds
are greatlydissimilar, left cheliped larger. C.
latens has whitish colored claws and same
colored dactylus of 2nd and 3rdpereopods.
Minimal setae, mainly on smaller right
cheliped, setae omitted on the outer margin of
palm. Setal brush are omitted on pereopod 3 and
has dark tipped pereopod 2 and 3. It has a
terminal cornea and slender moderate-
lengthened antennular peduncles.
Genus Clibanarius Dana 1852
Clibanarius cruentatus H. Milne Edwards,
1848
Clibanarius cruentatus are
observed to have white blotches and spots on
cephalothorax and telson but mostly towards
the ends of its slender and cylindrical
pereopods which have black tips. The
specimen used measures up to 40 mm and
shield length (SL) up to 21 mm. Covered with
limited small setae on its brown to purplish
pereopods and shield, the chelipeds are equal
or subequal and with no transverse striations
on pereopods 1-3, characteristics of
Clibanarius species. Dark corneas not
occupying more than one-third of slim
eyestalks, left longer than the right.
Clibanarius ransoni Forest, 1953
Clibanarius ransoni observed
specimen has a total length (TL) of 32 mm and
shield length (SL) up to 17 mm, has equal or
subequal chelipeds, pereopod 1-3 with black
tips, white colored dactylus of 2nd and
3rdpereopod and white purplish color on claw
with white spots on pereopod 1 with numerous
small setae. It has a dark purple band between
carpus and merus of pereopod 1, brownish to
orangey colored eyestalk, short antennula
peducles, and chelipeds. C. ransoni has a
terminal cornea and owns minimal setae,
found mainly on spinous chelipeds, omitted on
lateral part of 2nd and 3rd pereopod, white to
purplish shield with setae on sides.
Clibanarius virescens Krauss, 1843
Clibanarius virescens can be
easily distinguished with the presence of
yellow dactylus with pale brown mark. They
5
are known to have a pale brown colored
cephalothorax, pereopods covered with scarce
number of small setaes, pereopods darker on 2
and 3 and pereopod 1 being the darkest. The
pereopod 1-3 are all with black tips. Pereopod
1 with white spots frequently and bigger on its
merus and has an equal or subequalspinous
chelipeds. It also displays strong dark brown
bands on ends of its prodopus of pereopod 2
and 3 and alike colored marking on both ends
of its merus and on the shield. Its cornea does
not occupy 1/3 of its slender parallel eyestalks.
Material examined has a total length (TL) of 50
mm and shield length (SL) up to 30 mm.
Genus Dardanus Paul’son 1875
Dardanus deformis (H. Milne Edwards, 1836)
Dardanus deformis species have
setae predominantly on right cheliped and
partially omitted on the palm of the left.
Chelipeds are greatly unequal and dissimilar
which do not possess stridulating apparatus.
Spinous left chelipeds are evidently larger and
broader than right chelipeds but both possess
darkened tips. Rostrums are broadly rounded
or obsolete distinctive characteristics of
Dardanus species. Propodus of the left third
pereopod are considerably broader than the
right third pereopod (Malay et al., 2018).
Antennal peduncles are slender and cornea
occupies one-third of its white- and red-
banded ocular peduncles.
Dardanus megistos J. F. W. Herbst, 1804
Material Examined has up to total
length (TL) of 75 mm, shield length (SL)
measures up to 40 mm. specimen is
exceedingly covered with lengthy red setae
along its pereopods and chelipeds, minimal
undersized setae on its shield, telson, and parts
of the abdomen. They are mostly characterized
with their orange coloration with presence of
white spots. Chelipeds are greatly unequal and
is similar, a characteristic of Diogenidae
family, its left cheliped bigger and longer than
right but both have dark tipsd. Antennular
peduncles slender, not truncated and green
terminal cornea not does not occupy one-third
of its red ocular peduncles.
Dardanus lagopodes (Forskål 1775)
Specimen measured 68 mm in
total length (TL) and shield length (SL) up to
40 mm. Recognized by having red bands on
the shield, on carpus and merus of the second
and third pereopods. Mainly covered with
setae particularly on prodopus and dactylus of
pereopods, its left cheliped is somewhat longer
and larger than the right. Often mistaken as its
closely related species, Dardanus
sanguinolentos, D. lagopodesdo not have a
deep sulcus on thelateral face of left third
pereopod (Malay et al., 2018). Rostrum is
almost obsolete and antennal flagellum are
slender with darkened tips on it pereopods.
Genus Pagurus, Dana 1851
Pagurus aff. moluccensis
Though identified as P.
moluccensis, the specimen exhibits red
markings on white background on its shield
and chelipeds on dorsal and mesial view while
P. moluccensis is characterized with similar
pattern but with black colorations rather than
red (Haig, 1988). As the only Pagurus species
that was collected, the collected specimen
displays a pale colored shield and possess
equal or subequal chelipeds, right slightly
broader and longer than left. The short
eyestalk is mixed with orange and white color
and has a black cornea. It has slender orange
and white colored antennal peduncles slightly
longer than right 3nd pereopod.
A list of species arranged by their
taxa is given in table 1. There were 3 families
recorded in Verde Island namely are:
Coenibitidae, Diogenidae, and Paguridae.
Family Diogenidae contained more species. A
total of 12 species of hermit crab observed in
Verde Island, 8 of these were under the Family
Diogenidae; 3 for Coenobitidae; and only 1
species were observed in Paguridae.
In table 2, the summary of
biogeographic distribution of intertidal hermit
6
crabs was presented. Records of hermit crabs
from the Philippines were based from the
research of Malay et al. (2018). Clibanarius
virescens & C. cruentatus were previously
reported in Cebu and in Bohol (Cassidy &
McLaughlin, 1997; Malay et al, 2018) though
it wasn’t reported whether they have high
abundance as well. Diogenes sp. was recorded
in Cebu, Bohol, Balicasag, Palawan, Panglao,
Mindoro, and Puerto Galera (Estampador,
1959; Malay et al 2018; Malay and Palauy
2009; Rahayu et al, 2014). Similarly, although
coast of different Islands in Philippines have
been extensively sampled by the above
mentioned researchers, the species Cl. ransoni
were not found.
A total of 594 hermit crabs
(Diogenidae; Paguridae and Coenobitidae),
twelve species in 5 genera, were collected for
two sampling period during low tide in Verde
Island. The family Diogenidae contained more
species but family Coenobitidae had more
individuals. Coenobita rugosus H. Milne
Edwards, 1837 was the most abundant species,
with 285 specimens, followed by Clibanarius
virescens Krauss, 1843 with 132
specimens, Coenobita cavipes Stimpson, 1858
with 88 specimens (Table 3). Results of the
rapid record of common intertidal hermit crabs
demonstrate that the relative abundance of
hermit crabs varied from their substrate.
Among these Clibanarius virescens were the
most abundant in both sampling period. High
abundance level of family Diogenidae was
observed in rocky substrate. Intertidal hermit
crabs were most abundant during dry season.
Discussion
Family Diogenidae are known as
left- handed hermit crab for their left cheliped
is bigger than their right. They are the second
largest family of marine hermit crabs. Family
Diogenidae is the most species-rich group in
shallow waters, but so far in this data only 8
species in 3 genera is represented: Calcinus,
Clibanarius, and Dardanus. Calcinus is a
tropical to subtropical genus distributed from
supratidal to intertidal habitats all the way
down to deeper slopes. Most species are
associated with coral reefs. They are typically
very colourful, and live colouration is
important for species identification (e.g.,
Poupin& McLaughlin, 1998; Poupin&
Lemaitre, 2003). The species of Clibanarius
are restricted to the upper intertidal zone.
Members of this genus tend to be tolerant of
low salinities, often inhabiting estuaries and
mangrove areas. The genus Dardanus is
mostly large hermit crabs that are found in
shallow waters to deeper slopes; in tropical,
subtropical, to temperate latitude (Nakasone,
1988; Sato et al., 2013; Hamasaki et al., 2015).
Forty-four valid species have been described
(McLaughlin et al., 2010; Rahayu, 2010), of
which 30 are found in the Indo West Pacific
and 12 were previously reported from the
Philippines; 3 species are recorded in Verde
Island, Batangas City, Batangas, Philippines.
Pagurus Fabricius, 1775 is the most species-
rich hermit crab genus in the family Paguridae,
represented by about 170 species worldwide
(McLaughlin et al., 2010), but, only (1) one
species were recorded in the area, Pagurus aff.
moluccensis. The usual color of Pagurus
moluccensis are black and white, however our
collected specimen has red and white color,
either because of the preservation or this
species has really red and white color. Further
study is needed for this specimen.
Family Coenobitidae are known to
be terrestrial hermit crabs, which includes only
two genera; the land hermit crab genus
Coenobita Latreille, 1829 with 16 species,
only 3 species of this genera were recorded in
Verde Island. In table 2, consistent finding of
Dardanus sp in shallower water in different
island on Philippines by Estampador( 1959),
Malay et al. (2018), Malay and Palauy (2009),
& Rahayu et al. (2014), and in this research
indicate that these species are consistent
representative of shallow water which were
also observed in Fransozo et al. (2007) work.
Estampador’s list of decapodes was from
reports of previously surveys (US Exploring
Expidition (1852) & Siboga Expedition
(1920)) which shows that these species reigns
the Philippines century ago. Clibanarius
virescens & C. cruentatus hermit crabs are
7
reported to be sympatric in one specific area.
Turra & Denadai (2002) study revealed
Clibanarius sp. tends to be dependent on the
presence of one species on the presence of
another (Calcinus sp). This study could
support why these species found in same
places. The species Pagurus aff. moluccensis
was previously found in Panglao Island in
Bohol (Komai & Rahayu, 2014; Malay et al,
2018). This report extends the known
distribution substantially northward.
In table 3, family Coenobitadae
were separated to Family Diogenidae
Paguridae, because family Coenobitadae were
collected near the wells, underneath the
coconut tree, coastal areas soil because these
species can penetrate long distances. They can
live in both land and beaches where there is a
presence of fresh water. Coenibita sp were
mostly found in soils, roads or near wells, or
under the coconut trees. Some residents use
coconut latex as bait for hermit crabs to catch
them. These terrestrial anomurans became
catholic feeders and eat fallen fruits and seeds,
mangrove propagules, a wide variety of
otherplant material, strand line detritus, animal
faeces and animal carcases varying from small
invertebrates to fish, giant tortoises, birds,
goats and donkeys (Grubb, 1971; de Wilde,
1973; Vannini, 1976; Barnes, 1997).
Family Diogenidae and Paguridae
are known to be the intertidal hermit crab.
They are mostly collected in littoral to
sublittoral areas. In the study done by Synder-
Conn (1981) regarding adaptive significance
of clustering in the hermit crab which were
observed during the sampling was because of
the predictable and unpredictable
environmental changes which gives multiple
advantages to them. These include facilitation
of shell exchange, hermit crab tends to
withdraw into their shells to varying degrees
and thereby control the rate of evaporation and
desiccation; improve protection from strong
currents, desiccation, and predator avoidance
(Brock and Riffenburgh, 1960; Vine, 1971).
High abundance level of family Diogenidae
was observed in rocky substrate. High
abundance in rocky substrate was also
observed in other studies (Fransozo et al.,
2007). According to them, these site provide
wide variety of microhabitats, aside from
providing them protection against strong
waves and other factors, it can also serve as
habitat of other organisms which became food
for Paguroideans. In this research, a small part
of Verde Island was covered as a sampling site
(60m latitudinal and 10m longtidunal) in both
Sampalocan and Mahabang Buhangin.
Mahabang Buhangin has sandy course during
high tide but during low tide, coralline
substrate emerge revealing different marine
macroinvertebrate species. In Sampalocan,
where abiotic factors can be observed, it is
where boats from/to port stop and where lot of
residents does their recreational activities.
The value of pH ranges observed
in Verde Island range from 8.52 to 8.59 which
is the normal pH level for sea water.
Researchers can’t tell whether there are
changes in their behavior in rising and lowing
level of pH since the range of collected pH
values is small. Changes in temperature vary
the abundance of intertidal hermit crabs based
on their behaviours.
Conclusion
These findings shows that Verde
Island is an important area when it comes to
maintaining and establishing of Paguroidean
species despite of it being constantly by local
fishermen. However, as time pass by, Verde
Island has become hot spot for tourist, local or
international because of this several threats are
imposed to this marine biodiversity and
special attention is needed in conserving this
area in order to maintain Paguriodean species
richness in this area.
Recommendation
From the result, the researchers
recommend the following: (1) Make the
sampling site broader and deeper, since, a lot
of hermit crabs also occur in depth 10-40 m,
SCUBA diving technique is recommended to
do this (2) Include environmental factors such
as sediment samples, salinity, and dissolved
oxygen (DO) for higher accuracy of data; and
(3) Check mollusk shells availability and
8
 utilization pattern of hermit crabs present in
the sampling site to distinguish their
ecological and behavioral pattern concerning
their shell utilization.
Acknowledgement. We would like to express our
deepest gratitude to our Adviser, Prof. Ruthela P.
Payawal for supporting, guiding, and being with us
through the entire study and managing our stay in
Batangas as well as for helping us during our defense.
We also would like to thank the members of the panel,
Asso. Prof. Armin S. Coronado, Ms. Sarah Jane M.
Balana and Mr. Alejandro Jose C. Reboa for guiding us
through their suggestions and constructive remarks to
improve this research study. We would also like to
acknowledge Dr. Maria Celia Malay and Dr. Akira
Sakura for helping us identifying the hermit crabs
collected.
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