Shallow-water Sea Cucumbers (Echinodermata: Holothuroidea) in Sarangani Bay, Mindanao, Philippines with Notes on Their Relative Abundance

Abstract

The Philippines is known for its high diversity of sea cucumbers; however, there is little done on sea cucumber diversity in the southern Philippines. To augment this paucity of information, a rapid survey of the shallow-water sea cucumbers was carried out in three sites in Sarangani Bay by using the transect method for four months in both the seagrass and coral reef habitats. We recorded 21 species of sea cucumbers – 17 of these species were found in the seagrass beds and 10 species in the coral reef areas. The three most abundant species observed were the Holothuria scabra (29.2%), Bohadschia marmorata (21.2%), and Actinopyga echinites (17.6%) (n=1,969). Aside from some economically important sea cucumbers observed, we also noted rare species such as the Holothuria immobilis and Actinopyga capillata, which only have a few records in the Philippines. We are also the first to document Holothuria isuga in the Philippines, which is previously known only from five localities. The results offer opportunities for sea cucumber studies in the southern Philippines, especially on the biology and ecology of the rare and newly recorded species.

Full text

Key words: Actinopyga capillata, Holothuria immobilis, Holothuria isuga, holothurian, new
species record, Sarangani
Shallow-water Sea Cucumbers (Echinodermata:
Holothuroidea) in Sarangani Bay, Mindanao,
Philippines with Notes on Their Relative Abundance
1Animal Biology Division, Institute of Biological Sciences,
University of the Philippines Los Baños, College, Laguna 4031 Philippines
2Department of Marine Biology, College of Fisheries, Mindanao State University,
General Santos City 9500 Philippines
*Corresponding author: pitogo.kiermitchel@gmail.com
Kier Mitchel E. Pitogo1*, Jennelyn P. Sumin2, and Ariel T. Ortiz2
The Philippines is known for its high diversity of sea cucumbers; however, there is little done on
sea cucumber diversity in the southern Philippines. To augment this paucity of information, a
rapid survey of the shallow-water sea cucumbers was carried out in three sites in Sarangani Bay
by using the transect method for four months in both the seagrass and coral reef habitats. We
recorded 21 species of sea cucumbers – 17 of these species were found in the seagrass beds and
10 species in the coral reef areas. The three most abundant species observed were the Holothuria
scabra (29.2%), Bohadschia marmorata (21.2%), and Actinopyga echinites (17.6%) (n=1,969).
Aside from some economically important sea cucumbers observed, we also noted rare species
such as the Holothuria immobilis and Actinopyga capillata, which only have a few records in
the Philippines. We are also the first to document Holothuria isuga in the Philippines, which
is previously known only from five localities. The results offer opportunities for sea cucumber
studies in the southern Philippines, especially on the biology and ecology of the rare and newly
recorded species.
Philippine Journal of Science
147 (3): 453-461, September 2018
ISSN 0031 - 7683
Date Received: 21 Feb 2018
INTRODUCTION
The Philippines harbors a high diversity of sea
cucumber species, which is threatened by overfishing
and habitat loss (Choo 2008). Of the over 170 species
identified in the country (Olavides et al. 2010), 25 to
41 of these – mostly from the members of the families
Holothuriidae and Stichopodidae – have commercial
value (Gamboa et al. 2004; Olavides et al. 2010;
Jontila et al. 2014). Being a major exporter of trepang
or dried sea cucumbers worldwide (Akamine 2005),
the Philippines has been identified as a sea cucumber
hotspot in Asia (Choo 2008). This demand may lead to
overharvesting of sea cucumber resources, which may
pose a great risk to their populations. The declining
numbers of sea cucumbers in intertidal areas have
serious consequences in the ecosystem such as habitat
structure alteration, as they are the seas’ bioturbators
and recyclers (Wolkenhauer et al. 2010). Moreover,
sea cucumber species contain compounds that have
medicinal properties that are of great importance to
future research (Bordbar et al. 2011).
Most of the commercially-important sea cucumbers that
inhabit the shallow waters have been fished with increasing
intensity (Wolkenhauer et al. 2010). Shallow-water sea
453

cucumbers, especially, are vulnerable to harvesting because of
their high abundance and easy access to their habitats (Purcell
et al. 2013). With this, it is imperative that baseline research
(e.g., inventory studies) is needed to support science-based
conservation and management measures as the demand for
sea cucumbers increase. While there were several diversity
studies of sea cucumbers conducted in the Philippines, none
has been conducted in Sarangani Bay. Anecdotal evidence
from local interviews shows that overharvesting caused a
reduction of a high-valued Holothuria scabra in Macatimbol,
Glan, Sarangani Province, as buyer boats from Davao once
frequent the area to buy pails of H. scabra from the locals
(Pitogo et al. 2016). If harvesting continues unregulated
and if preferences shift to other species of similar or lower
value, local sea cucumber populations may collapse leading
to serious environmental repercussions. Thus, this study
was carried out to provide a preliminary inventory of the
sea cucumber species found in the shallow-water areas in
the bay with information on their relative abundance. The
results of this study may provide helpful insights in creating
local policies regarding the management of the sea cucumber
resources in the area.
MATERIALS AND METHODS
Sarangani Bay is in the southeastern Mindanao,
Philippines between 5º33’25’’- 6º6’15’’N and 124º22’45’’-
125º19’45’’E. There were three stations in Sarangani Bay
selected in the study: Sitio Linao and Sitio S’nalang,
Figure 1. The map of Sarangani Bay showing the three study sites.
Maasim; and Sitio Macatimbol, Glan. The stations
were chosen based on the reported high abundance of
sea cucumbers. All stations have seagrass beds covered
mainly by Cymodocea sp., Halodule sp., and Halophila
sp. on a sandy and silty substrate. The Macatimbol station,
however, had a patchier distribution of seagrass on a
sandy and rubble substrate. Of the three sites, the Sitio
Linao station had the most minimal wave action because
of its placement in a cove. It also had the siltiest substrate
dominated by the seagrass Enhalus acoroides. The reef
formation in all three sites was dominated by massive
and sub-massive corals creating pools of water during the
lowest ebb. This leaves patches of sand and rubble usually
exposed that were used as trails to minimize disturbance.
At each station, six transects measuring 4 m x 50 m were
laid perpendicular to the shore with a 10-meter interval.
Three transects were laid in the seagrass bed and the other
three were laid in the coral reef area. We completed 24
night samplings during the lowest ebbing of the month
from Oct 2014 to Jan 2015. Three investigators, with one
local guide adept at collecting sea cucumbers, collected
all sea cucumbers found within the 200 m2 transects by
reef walking at around 22:00-23:00 to 02:00-03:00. For
some species, we used our bare feet to locate individuals
burrowed into the sand (a method used by some locals).
One specimen for each species was collected for
photo documentation. The species were identified by
morphology using the book Commercially Important
Sea Cucumbers of the World by Purcell and co-authors
(2012). The photographs of identified and unidentified
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
Mindanao, Philippines
Philippine Journal of Science
Vol. 147 No. 3, September 2018
454

species were sent to Dr. Gustav Paulay and Dr. Francóis
Michonneau of the Invertebrate Zoology Division at the
Florida Museum of Natural History for validation and
identification.
RESULTS AND DISCUSSION
We recorded 21 species of shallow-water sea cucumbers
belonging to families Holothuriidae and Stichopodidae,
17 of these species were found in the seagrass beds and 10
species in the coral reef areas. This is relatively lower than
the other species diversity studies in the Philippines (see dela
Cruz et al. 2015). This low number recorded might be due
to the shallowness of the area assessed or due to our limited
number of sampling stations and the total area surveyed.
Of the 1,645 individuals counted in the seagrass beds,
the species Holothuria scabra (34.9%), Bohadschia
marmorata (25.3%), and Actinopyga echinites (20.8%)
were the most commonly observed. Species belonging
to these three genera tend to be the most common in the
shallow seagrass beds, similar with the observations of
Jontila and co-authors (2017) and Olavides and co-authors
(2010). Specifically, species belonging to Holothuria
accounted for 52.1% of the sea cucumbers found in
the seagrass beds, while Actinopyga and Bohadschia
accounted for 21.2% and 26.1%, respectively.
Holothuria fuscocinerea, the most dominant in the coral
reef areas, accounted for 68.2% of the 324 individuals
found. Individuals of the species hid most of their
body under rocks, leaving only a small portion outside
the crevice. When disturbed, they start writhing and
injecting numerous thick Cuvierian tubules, similar with
a predatory escape behavior of H. fuscocinerea observed
in the laboratory by Morton (1991). We suspect that H.
fuscocinerea is sensitive to light as retraction was observed
upon flashing light to individual organisms. This might be
because of its strict nocturnal behavior (Kerr et al. 2006;
Purcell et al. 2012; Jontila et al. 2017). Despite being a
common species, ecological studies on H. fuscocinerea
is very little to support our observations.
Fewer sea cucumber species was recorded in the coral
reef areas. We had to move at a slower pace in the reef
areas while we were sampling to avoid causing damage to
Table 1. The initial list of sea cucumber species found in the shallow waters of Sarangani Bay.
Species Common Name Local Name Habitata
1Actinopyga capillata Hairy sea cucumber SG
2Actinopyga echinites Deep-water redsh Bat-alaw/Hud-hud SG, C
3Actinopya lecanora Stonesh Batseror SG
4Actinopyga sp.bSG
5Bohadschia argus Leopardsh Bangkungan C
6 Bohadschia koellikeri Mottled sea cucumber Tagukan SG,C
7Bohadschia marmorata Brownspotted sandsh Tagukan SG
8Bohadschia vitiensis Brown sandsh Tagukan SG,C
9Holothuria albiventer Marten’s sea cucumber SG
10 Holothuria atra Lollysh Bat-uwak SG,C
11 Holothuria fuscocinerea Variegated sea cucumber Gulay-gulay/Libud-buto SG,C
12 Holothuria hilla Tigertail sea cucumber Pula-pula/Iring-iring C
13 Holothuria immobilisbLibud-buto C
14 Holothuria isugabSG
15 Holothuria leucospilota White threadsh Patola SG,C
16 Holothuria notabilis SG
17 Holothuria rigida Rigid sea cucumber Bantunan SG
18 Holothuria scabra Sandsh Putian SG
19 Holothuria sp.bSG
20 Stichopus horrens Warty sea cucumber Hanginan C
21 Stichopus a. horrensbWarty sea cucumber Hanginan SG
a SG= seagrass; C= coral reefs
bnot included in the list of the shallow-water sea cucumbers compiled by dela Cruz and co-authors (2015)
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
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Philippine Journal of Science
Vol. 147 No. 3, September 2018
455

the corals. There were instances that sea cucumbers have
already retracted to crevices even before we approached
them, probably because of the light emitted from our lamp.
This factor, together with the fact that coral reef areas have
a lot of nooks and crannies where sea cucumbers could
hide, might have contributed to the low number of species
recorded in this habitat. The rock crevices were favorite
hiding spots of slender-bodied sea cucumbers such as the
H. fuscocinerea (Purcell et al. 2012). Hiding in crevices is
one of the most common self-protecting behaviors of sea
cucumbers aside from burrowing (Purcell 2010). Unlike in
coral reef areas, spotting sea cucumbers was easier in the
seagrass beds as they were usually found exposed, except
during extreme ebb conditions. Only a few of the recorded
species in the seagrass beds bury their body underground
such as the Holothuria rigida, Holothuria albiventer, and
Holothuria notabilis, which made these species difficult
to spot. This burrowing behavior of sea cucumbers,
however, plays a critical role in mixing the sediment layers
in the intertidal ecosystem. The commonly observed H.
scabra that moves in and out of the sub-surface sediments
are probably the most important bioturbators in this
environment (Wolkenhauer et al. 2010). Sea cucumbers
also increase the nutrient levels in the water column (Grall
& Chauvaud 2002) and, as deposit feeders, clean the
sediments by lowering its organic matter upon defecation
(MacTavish et al. 2012). Collectively, holothurians play
a significant role in the intertidal ecosystems by helping
increase the productivity of these environments (Purcell
Figure 2. The pooled relative abundance of shallow-water sea cucumbers surveyed in the 1,200 m2
area of both coral reefs and seagrass beds in Sarangani Bay, Philippines from Oct 2014 to
Jan 2015 (n=1,969).
et al. 2013). Uncontrolled harvesting of holothurians that
perform this function would eventually lead the collapse
of intertidal ecosystems (Wolkenhauer et al. 2010).
Of the 1,969 sea cucumber individuals recorded, H.
scabra had the highest occurrence (29.2%), followed
by B. marmorata (21.2%) and A. echinites (17.6%). H.
scabra prefer seagrass beds, which they rely heavily on
for settling cues and during their early life stages (Mercier
et al. 2000). In all three sites, H. scabra had the highest
occurrence in Sitio Linao where wave action was minimal
and organic matter concentration was high (Pitogo et
al. 2016). In 2013, however, sea cucumber pens were
established in Sitio Linao to rear hatchery-produced H.
scabra. The individuals were eventually released after
the project, which might have influenced the increase
in H. scabra abundance in succeeding years. Locals in
Sarangani Province collect H. scabra, preferably during
a new moon phase, where larger individuals occur (Pitogo
et al. 2016). They are high-value species, which are
harvested and exploited leading to the depletion of their
natural population (Akamine 2005). Other commonly
occurring species recorded in the seagrass beds were B.
marmorata and A. echinites, low-value and medium-value
species, respectively (Olavides et al. 2010).
Despite the limited number of our sampling stations and
the total area surveyed, a few notable species were found.
One of our interesting find was the species Holothuria
isuga. It is a new record for the Philippines as the reported
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
Mindanao, Philippines
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Vol. 147 No. 3, September 2018
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Figure 3. Photos of the four relatively rare sea cucumber species recorded: (A) Stichopus aff. horrens, (B)
Actinopyga capillata, (C) Holothuria immobilis, and (D) Holothuria isuga.
ranges for this species are Japan, Australia, Tanzania,
and Sri Lanka (IUCN 2013a). Very few studies have
mentioned H. isuga: in Zanzibar, Tanzania (Eriksson et
al. 2010); in Sri Lanka (Dissanayake & Stefansson 2010);
and another in New Caledonia (Purcell et al. 2009). It was
described by Purcell and co-authors (2009) as “rusty-
brown, with large papillae and buries most of its soft
body in sediments”. We only found one H. isuga partly
buried in silt near the mangrove area at night. The time
of our observation was not similar with Dissanayake and
Stefansson (2010), who observed H. isuga at daytime. The
activity pattern of H. isuga is not yet fully known since
it is rarely seen, at least in some areas. The H. isuga is
characterized by its soft body covered with black papillae
with a narrow orange streak along the dorsum and on the
ventrum, with a high amount of similar color around its
anal portion (see Figure 4). It also writhes and distorts
its body when touched or handled. Divers do not often
visit sandy and silty habitats where the H. isuga lives
buried that is why this species is rarely cited in literature
(Dr. Francóis Michonneau, pers. comm., 04 May 2016).
Interestingly, Purcell and co-authors (2009) and Eriksson
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
Mindanao, Philippines
Philippine Journal of Science
Vol. 147 No. 3, September 2018
457

and co-authors (2010) collected them in lagoon reef areas
while we saw our specimen in the silty substrate near
a mangrove area. Our observation supports the habitat
association of this species to silty environment, as the
few previous studies only observed them in the reef areas.
Despite their current rarity, their burrowing behavior
still makes them an important bioturbator in the seagrass
beds (Purcell et al. 2016). Unfortunately, they are being
harvested in some parts of their distribution (Eriksson et
al. 2010; Eriksson et al. 2012). Previous records of these
species suggest that this species occurs throughout the
western Pacific Ocean and in the northern Indian Ocean
(IUCN 2013a). But with this new record, H. isuga is
maybe more widespread than its current reported range.
The species Holothuria immobilis is also rarely cited in sea
cucumber diversity studies and was only collected twice
in the study of dela Cruz and co-authors (2015) in Samar
and Leyte. It was noted by Cherbonnier (1967) that the
species can be found in the Philippines as described by
Semper in 1868, but generally has been lost in synonymy.
Information regarding the population status, habitat,
and ecology of this species is very little (IUCN 2013b).
Although Clark reported the species in Torres Strait
in 1921, it remained an invalid species and had been
forgotten until it was rediscovered by Dr. Gustav Paulay
and his team and confirmed its validity with genetic data
(Dr. Francóis Michonneau, pers. comm., 28 Mar 2016).
Ten H. immobilis were found hiding in rock crevices in
the coral reefs in Sitio Linao with two different coloration
patterns: a dark brown and a light brown with dark
blotches on its dorsum (see Figure 4). The dark morph
of H. immobilis is quite similar with the H. fuscocinerea.
However, it is easily distinguished by its blue-tipped
papillae, as opposed to the dark white-ringed papillae of
H. fuscocinerea (see Figure 4). The H. immobilis is known
in two localities in Sarangani: Sitio Linao, Maasim (this
study) and Sitio Pananggalon, Malapatan (Gino 2016).
Gino (2016) also found H. immobilis along the reef flats
hiding in rock crevices. Interestingly, all H. immobilis
were observed during a new moon phase where it is
relatively darker. This may suggest a strict nocturnal
behavior of this organism like the A. capillata (Rowe &
Massin 2006).
A strictly nocturnal species of sea cucumber, A. capillata,
was recorded five times in the seagrass beds of Sitio
S’nalang only during a new moon phase. It is characterized
by its numerous hair-like papillae and mottled light to dark
orange and white on its body (see Figures 3 & 4). It was
only recently described by Rowe and Massin (2006) from
a holotype specimen collected in La Réunion. We found
all the individuals submerged in both sandy and rubble
substrates in the seagrass area, which probably is a new
habitat record for this species. The holotype and paratype
specimens were collected in the inner reef and reef crests
Figure 4. (A) Holothuria isuga submerged in water highlighting its rusty brown papillae; (B) the blue-
tipped papillae of Holothuria immobilis (above) and the dark white-ringed papillae of Holothuria
fuscocinerea (below); (C) the appearance of the nocturnal Actinopyga capillata submerged in
water in a seagrass bed in Sitio S’nalang, Maasim, Sarangani Province.
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
Mindanao, Philippines
Philippine Journal of Science
Vol. 147 No. 3, September 2018
458

similar with where Ducarme (2017), Lampe-Ramdoo and
co-authors (2014), and Conand and co-authors (2010)
observed them. Our observations, together with those
of Kerr and co-authors (2006), extend the distribution
of A. capillata from the seagrass beds and algal beds to
the reef crest. Interestingly, the present distribution of A.
capillata is disjunct with one in Mayotte to Mauritius and
the other in the Philippines (Ducarme 2017). There used
to be one record of A. capillata in the Philippines, but a
few inventory studies mentioned the species though only
in few numbers (Kerr et al. 2006; dela Cruz et al. 2015).
It seems that based on records, the A. capillata’s current
distribution in the Philippines is in the central (Kerr et al.
2006; dela Cruz et al. 2015; Ducarme 2017) and southern
(Ducarme 2017, this study) part. In the study of Kerr and
co-authors (2006), they recorded A. capillata in the central
Philippines but they misidentified it (named it Holothuria
sp.). While in a recent study of dela Cruz and co-authors
(2015), 15 individuals were found in the shallow waters of
Samar and Leyte, but they did not include field observations
that could have been important information on its habitat
associations. Most A. capillata data are only accounts
with limited information on its biology and ecology. Our
observations concur with the original description of Rowe
and Massin (2006) that this species is nocturnal and occur
in shallow-water habitats. We suspect that the Philippine
population may also be cryptic and strictly nocturnal, like
in some of its possible distributional range (Ducarme 2017),
due to its rare appearance in the sea cucumber diversity
studies in the country.
It is worth noting that a Stichopus aff. horrens was found
in a silty substrate in a patch of seagrass near a mangrove
area. The Stichopodids usually live exposed on sand and
rubble in the reef areas and do not usually inhabit silty
environments (Purcell et al. 2012). This reddish variant
was not reported in previous literature, but something
similar was once collected by Dr. Francis Michonneau
in Papua New Guinea (Dr. Francóis Michonneau, pers.
comm., 29 March 2016) (see Figure 3). However, further
validation through ossicles and molecular data is needed
to validate our identification.
Our results only provide a glimpse of the sea cucumber
diversity in Sarangani Bay. Despite the limited sampling
stations and the total area surveyed, we still managed
to record 1,969 individuals belonging to 21 species of
economic and ecological importance. We did not have
enough information on the biology and ecology of the
species since we only collected abundance data and
took photographs. Our results, however, highlights
the importance of Sarangani Bay in sea cucumber
research, especially that anecdotal evidence shows that
overharvesting and direct exploitation threatens the
local sea cucumber populations in the area. Moreover,
our results warrant species-specific studies to provide
significant information for the relatively rare species
observed, which could provide helpful data for the
formulation of conservation and management plans
in the area.
ACKNOWLEDGMENTS
Gratitude is given to Dr. Gustav Paulay and Dr. Francóis
Michonneau of the Invertebrate Zoology Division at
the Florida Museum of Natural History for validating
and identifying sea cucumber species, and for the
useful information they shared that helped in finishing
the manuscript. We thank the Sarangani Bay Protected
Seascape (SBPS) Protected Management Board for
allowing us to conduct research in Sarangani Bay
(Certification No. SBPS-014-009). We would also like
to extend our thanks to the two anonymous reviewers for
their helpful comments and suggestions.
REFERENCES
AKAMINE J. 2005. Role of the trepang traders in the
depleting resource management: A Philippine case.
In: Kishigami N, Savelle JM. Indigenous use and
management of marine resources. Senri Ethnological
Studies 67: 259-278.
BORDBAR S, ANWAR F, SAARI N. 2011. High-value
components and bioactives from sea cucumbers for
functional foods—A review. Marine Drugs 9(10):
1761-1805.
CHERBONNIER G. 1967. Deuxieme contribution a
l'etude des Holothuries de La Mer Rouge collectees
par des Israeliens. Isreal South Red Sea Expedition,
1962, Reports No. 26.
CHOO PS. 2008. The Philippines: a hotspot of sea
cucumber fisheries in Asia. In: Toral-Granda V,
Lovatelli A, Vasconcellos M (eds). Sea cucumbers:
A global review of fisheries and trade. FAO Fisheries
and Aquaculture Technical Paper. No. 516. Rome,
FAO. p. 119-140.
CLARK HL. 1921. The echinoderm fauna of Torres Strait:
Its competition and its origin (No. 214). Carnegie
Institution of Washington, Washington, D.C.
CONAND C, MICHONNEAU F, PAULAY G,
BRUGGEMANN H. 2010. Diversity of the Holothuroid
Fauna (Echinodermata) at La Réunion (Western Indian
Ocean). Western Indian Ocean Journal of Marine
Science 9(2): 145-151.
DELA CRUZ MT, CABANSAG JB, GAJELAN-
SAMSON MBP, DIAZ FA, DIODOCO RJ. 2015.
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
Mindanao, Philippines
Philippine Journal of Science
Vol. 147 No. 3, September 2018
459

Diversity and Abundance of Shallow-water Sea
Cucumbers in Samar and Leyte, Philippines. Asian
Journal of Biodiversity 6(1).
DISSANAYAKE DCT, STEFANSSON G. 2010.
Abundance and distribution of commercial sea
cucumber species in the coastal waters of Sri
Lanka. Aquatic Living Resources 23(3): 303-313.
DUCARME F. 2017. Revision of the geographic range
of Actinopyga capillata Rowe & Massin 2006
(Echinodermata: Holothuroidea). SPC Beche-de-mer
Information Bulletin 37: 92-94.
ERIKSSON H, DE LA TORRE-CASTRO M, OLSSON P.
2012. Mobility, expansion and management of a multi-
species scuba diving fishery in East Africa. PLOS
one 7(4): e35504.
ERIKSSON BH, DE LA TORRE-CASTRO M., EKLÖF
J, JIDDAWA N. 2010. Resource degradation of the
sea cucumber fishery in Zanzibar, Tanzania: a need
for management reform. Aquatic Living Resources
23: 387-398.
GINO CF. 2016. Inter-dynamic relationship of sea
cucumbers (Holothuroidea) on selected mangrove,
seagrass, and coral reef areas in Sarangani Bay,
Philippines. [BS Thesis]. General Santos City, South
Cotabato: Mindanao State University. 55p. (Available
at the College of Fisheries, MSU-General Santos
Library)
GRALL J, CHAUVAUD L. 2002. Marine eutrophication
and benthos: the need for new approaches and
concepts. Global Change Biology 8(9): 813-830.
GAMBOA R, GOMEZ AL, NIEVALES MF. 2004. The
status of sea cucumber fishery and mariculture in the
Philippines. p. 69-78. In: Advances in sea cucumber
aquaculture and management. Lovatelli A, Conand
C, Purcell S, Uthicke S, Hamel JF, Mercier A. (eds).
2004. FAO Fisheries Technical Paper. No. 463. Rome,
FAO. 425p.
[IUCN] International Union for Conservation of Nature.
2013a. The IUCN Red List of Threatened Species.
Version 2013. Retrieved from http://www.iucnredlist.
org/details/summary/19710863/0
[IUCN] International Union for Conservation of Nature.
2013b. The IUCN Red List of Threatened Species.
Version 2013. Retrieved from http://www.iucnredlist.
org/details/full/19709103/0
JONTILA JBS, BALISCO RAT, BATIN GT. 2017.
Species composition, density and distribution of
sea cucumbers (Holothuroidea) at Arreceffi Island,
Honda Bay, Palawan, Philippines. SPC Beche-de-mer
Information Bulletin 37: 21-29.
JONTILA JBS, BALISCO RAT, MATILLANO JA.
2014. The sea cucumbers (Holothuroidea) of Palawan,
Philippines. AACL Bioflux 7(3): 194-206.
KERR AM, NETCHY K, GAWEL AM. 2006. Survey
of the shallow-water sea cucumbers of the Central
Philippines. University of Guam Marine Laboratory,
Technical Report No. 119, Jun 2006.
LAMPE-RAMDOO K, PILLAY RM, CONAND C. 2014.
An assessment of holothurian diversity, abundance and
distribution in the shallow lagoons of Mauritius. SPC
Beche-de-mer Information Bulletin 34: 17-24.
MACTAVISH T, STENTON-DOZEY J, VOPEL K,
SAVAGE C. 2012. Deposit-feeding sea cucumbers
enhance mineralization and nutrient cycling in
organically-enriched coastal sediments. PLoS ONE
7(11): e50031.
MERCIER A, BATTAGLENE SC, HAMEL JF. 2000.
Settlement preferences and early migration of the
tropical sea cucumber Holothuria scabra. Journal
of Experimental Marine Biology and Ecology 249:
89-110.
MORTON B. 1991. Aspects of predation by Tonnazonatum
(Prosobranchia: Tonnoidea) feeding on holothurians in
Hong Kong. Journal of Molluscan Studies 57(1): 11-19.
OLAVIDES RD, EDULLANTES CM, JUINIO-MEÑEZ
MA. 2010. Assessment of the sea cucumber resource
and fishery in the Bolinao-Anda reef system. Science
Diliman 22(2): 1-12.
PITOGO KM, SUMIN JP, ORTIZ AT. 2016. Effect of lunar
phases in the size distribution of Holothuria scabra on
intertidal areas in Sarangani Bay, Philippines. SPC
Beche-de-mer Information Bulletin 36: 48-53.
PURCELL SW, CONAND C, UTHICKE S, BYRNE
M. 2016. Ecological roles of exploited sea
cucumbers. Oceanography and Marine Biology: An
Annual Review 54: 367-386.
PURCELL SW, MERCIER A, CONAND C, HAMEL JF,
TORAL‐GRANDA MV, LOVATELLI A, UTHICKE
S. 2013. Sea cucumber fisheries: global analysis
of stocks, management measures and drivers of
overfishing. Fish and Fisheries 14(1): 34-59.
PURCELL SW, SAMYN Y, CONAND C. 2012.
Commercially important sea cucumbers of the world.
FAO Species Catalogue for Fishery Purposes. No. 6.
Rome, FAO. 150p.
PURCELL SW. 2010. Diel burying by the tropical sea
cucumber Holothuria scabra: effects of environmental
stimuli, handling and ontogeny. Marine biology 157(3):
663-671.
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
Mindanao, Philippines
Philippine Journal of Science
Vol. 147 No. 3, September 2018
460

PURCELL SW, GOSSUIN H, AGUDO NN. 2009. Status
and management of the sea cucumber fishery of la
Grande Terre, New Caledonia. Programme ZoNéCo.
WorldFish Center Studies and Reviews No. 1901. The
WorldFish Center, Penang, Malaysia.
ROWE FWE, MASSIN C. 2006. On a new species
of Actinopyga Bronn, 1860 (Echinodermata,
Holothuroidea) from the IndoWest Pacific. Zoosystema
28: 955-961.
WOLKENHAUER SM, UTHICKE S, BURRIDGE C,
SKEWES T, PITCHER R. 2010. The ecological role
of Holothuria scabra (Echinodermata: Holothuroidea)
within subtropical seagrass beds. Journal of the Marine
Biological Association of the United Kingdom 90(2):
215-223.
Pitogo et al.: Sea Cucumbers in Sarangani Bay,
Mindanao, Philippines
Philippine Journal of Science
Vol. 147 No. 3, September 2018
461