Symbiotically snapper shrimp Anchistus custoides Bruce, 1977 (Decapoda: Palaemonidae) inhabiting the mantle cavity of the pen shell (Bivalvia: Pinnidae)

eISSN: 2672-7226
© Penerbit UMT
Journal of Sustainability Science and Management
Volume 17 Number 10, October 2022: 136-148
SYMBIOTICALLY SNAPPER SHRIMP ANCHISTUS CUSTOIDES BRUCE, 1977
(DECAPODA: PALAEMONIDAE) INHABITING THE MANTLE CAVITY OF
THE PEN SHELL (BIVALVIA: PINNIDAE)
MOHD HANAFI IDRIS1
*, MUHAMMAD ATIF AWANG1
, ABU HENA MUSTAFA KAMAL1
,
AZIZ ARSHAD2
, HADI HAMLI3
, ABDULLA AL-ASIF3
, KUSTIAWAN TRI PURSETYO4
AND
ENDANG DEWI MASITHAH4
1
Faculty of Fisheries and Food Science, Universiti Malaysia Terengganu, 21030 Kuala Nerus, Terengganu, Malaysia.
2
Department of Aquaculture, Faculty of Agriculture, Universiti Putra Malaysia, 43400 UPM, Serdang, Selangor, Malaysia.
3
Department of Animal Science and Fisheries, Faculty of Agricultural Science and Forestry, Universiti Putra Malaysia
Bintulu Sarawak Campus, 97008 Bintulu, Sarawak, Malaysia. 4
Department of Marine, Faculty of Fisheries and Marine,
Universitas Airlangga, Kampus C Jalan Mulyorejo, Surabaya 60115, Jawa Timur, Indonesia.
*Corresponding author: hanafiidris@umt.edu.my
Submitted final draft: 21 June 2022 Accepted: 11 September 2022
Introduction
The relationship between decapods and
molluscs, known as symbiosis is a one-of-a-kind
chance to investigate the evolutionary methods
used by marine invertebrates (García-Ulloa et
al., 2019). Some invertebrates included small
decapods living in or on other organisms such
as marine molluscs, anemones, polychaetes
and echinoderms (Nizinski, 1989; Itani et al.,
2002; Baeza et al., 2011; Baeza et al., 2013).
They are an attractive method to determine the
development of connections between decapods
and molluscs since Pontonnid shrimps have
developed a number of adaptations to deal with
a symbiotic existence (Peiró & Mantelatto,
2011). These associations may arise when a
guest organism is looking for a haven from
its predators, investigates a new food niche or
uses its host as a place for reproduction (Baeza,
2010). However, because of the different
environmental conditions that impede direct
observation and identification of the specific
types of association, the relationship between
a pen shell and a snapper shrimp is frequently
not clearly defined (Radda & Milat, 2009;
Overstreet & Lotz, 2016).
Because many species are composed of
several different life stages, benthic habitats
provide interesting systems for testing the
working effect of different spatial scales
(Munguia, 2004). Previous research has used
the benthic creatures connected to the pen shells
Abstract: Snapper shrimp is a symbiotic organism usually hidden under the rocks,
sponges and pen shells in the seagrass and coral habitats. The relationship study within
snapper shrimp and pen shell was conducted from Merambong shoal, one of the biggest
seagrass beds in peninsular Malaysia. A total of 40 individual pen shells were collected
randomly and four species of pen shells were identified. 40 Anchistus custoides were found
inhabiting symbiotically in the mantle cavity of the pen shell as solitary males and females
and heterosexual pairs. Pen shell, Pinna bicolour and Atrina vexillum recorded the highest
average SH 217.79±53.15 mm, SV 2.62±1.36 dm3
and SH 164.10-224.78 mm with the SV
1.18±0.43 dm3
, respectively compared to the other species. The size of Anchistus custoides
ranged from 15.00 to 20.00 mm in length and it was determined to be female due to the
presence of eggs in the pleopods. The length of the cephalothorax and its length were
highly related (rs
=0.563, p≤0.01, N=40) and found wider in females. A little difference
in size between the left and right chela in males of identical length was noticed, although
the left chela is much bigger than the right. The significant relationship (rs
=0.450, p≤0.01,
N=40) between the pen shell length and shrimp (male-female) length revealed that the size
of the shell is important to be hosted the snapper shrimp in the shell cavity.
Keywords: Merambong shoal, pen shell, shell volume, snapper shrimp, symbiotic.
http://doi.org/10.46754/jssm.2022.10.010

PEN SHELLS EAST COAST MALAYSIA 137
Journal of Sustainability Science and Management Volume 17 Number 10, October 2022: 136-148
(Pinna bicolor) as a model system to analyse
the communities found in patchy environments
(Keough, 1984a; 1984b; Butler, 1987). There
have been reports of symbiosis between several
differentspeciesofpontoniidsandadiversegroup
of host taxa, which includes invertebrates such
as corals, jellyfish, sponges and molluscs (Lee
& Ko, 2011; Olliff, 2013; Dobson et al., 2014;
García-Ulloa et al., 2019; Chow et al., 2021).
One of the most well-researched symbiosis types
involves bivalve hosts colonised by tiny shrimp
(Baeza, 2008; Aucoin & Himmelman, 2010;
Baeza et al., 2013). Important indications used
to estimate the symbiotic relationship between a
host and a guest are decapods’ physical, sexual
and reproductive characteristics in proportion
to the size of the host (Baeza et al., 2015). In
tropical and subtropical climates where they are
found, the Pontoniinae family has more than
600 species that live within the first 100 meters
of depth (De Grave & Fransen, 2013). The shells
of many different kinds of marine bivalves,
particularly those belonging to the Pinnidae
family, served as homes for pontoniid shrimps
(Kennedy et al., 2001; Aucoin & Himmelman,
2010). Many marine bivalves, particularly those
belonging to the Pinnidae family are home to
pontoniid shrimps inside their shells (Richardson
et al., 1997; Kennedy et al., 2001; Rabaoui et
al., 2008; Aucoin & Himmelman, 2010).
In the region known as the Indo-Pacific, the
Pinnidae family may be found from southeastern
Africa through Melanesia and New Zealand, all
the way north to Japan and New South Wales
and New Zealand (Butler & Keough, 1981;
Poutiers, 1998). Additionally, pen shells may be
found in the seas of the Mediterranean and the
United States (Rosewater, 1961; 1982; Butler,
1987; Zavodnik et al., 1991; Butler et al., 1993;
Munguia, 2004). The pen shell is an endemic
species in the Mediterranean Sea (Katsanevakis,
2004) and the biggest Mediterranean bivalve
as well as one of the largest in the world,
reaching lengths of up to 120 cm (Zavodnik et
al., 1991). In contrast, the horse mussel, known
as Atrina zelandica is a huge pinnid bivalve
that feeds on suspended particles and is found
randomly spread around the coast of New
Zealand (Cummings et al., 1998). In Malaysia,
the distribution of pen shells (Pinna and Atrina)
was reported at Merambong shoal and Tanjung
Adang shoal (Johor), Bagan Panchor (Perak)
and Merchang Lagoon (Terengganu) (Idris et
al., 2008; 2009; Idris et al., 2012).
There is currently no extensive
documentation on the symbiosis between
the shrimp Anchistus custoides Bruce, 1977
(Decapoda: Palaemonidae) that occupy the
mantle chamber of the shell (Bivalvia: Pinnidae)
of Malaysian pen shells that have been reported.
In Malaysia, most pen-shell studies focus on
distribution and ecology (Idris et al., 2008;
2009). Most of the publications reported in
the Indo-Pacific, Mediterranean and American
regions a long time ago (Zavodnik, 1967; Butler
& Brewster, 1979; Butler & Keough, 1981;
Scheltema, 1983; Butler, 1987; Butler et al.,
1993; Šiletić & Peharda, 2003; Katsanevakis,
2004). This study focuses on discovering
symbiotic snapper shrimps in the pen shell
mantle cavity, which may eventually help to
know the characteristics and habitat information
of snapper shrimp living inside the pen shell
cavity in the seagrass bed of Malaysia.
Materials and Methods
During the time of low tide, a total of 40
different individuals of pen shells were collected
at random from the Merambong shoal in the
Johor Strait (N1o
19’ 55.62” E103o
35’ 57.75”)
(Figure 1). Merambong shoal is a sandy area
with Enhalus acoroides, Halophila ovalis,
Thalassia hemprichii and Halodule uninervis
growing in seagrass meadows. Specimens were
removed from their natural habitat using a hand
scoop and specimens were placed into a plastic
bag containing 10% formalin and labelled.
Specimens were transported to the laboratory
for further examination. In the laboratory, pen-
shell specimens were emptied into a washing
tray according to the labeled given and washed
in distilled water over a 0.5 mm sieve to avoid
the shrimp passing through with the running
water.

Mohd Hanafi Idris et al. 138
Pen shells adductor muscles were sacrificed
by having the back cut off to get the shrimp
occupying the shell. The shrimp were identified
based on the key features ofAnker and De Grave
(2016).Thenumberofshrimpsfoundoneachpen
shell was recorded and preserved in ethanol at a
70% concentration. The works of Richardson et
al. (1997) were adopted to identify the presence
of snapper shrimp inhabiting the pen shell.
Using a MITUTOYO digital vernier calliper
(±0.01 mm), the total body length (BL) (from
the tip of the rostrum to the tip of the telson),
cephalothorax length (CL) (from the tip of the
rostrum to the posterior end of the carapace) and
width of the widest part of the cephalothorax,
and length of the right and left chelae (from the
base of the dactyl and propodus to the tip of the
claw) of the second pereiopods of each shrimp
were measured. Morphometric relationships
between the characteristic of shrimp (BL and
CL) were analysed using Pearson’s correlation
(Bhujel, 2008).
Figure 1: The map displayed the sample regions, including (A) Merambong shoal, which is located off the
coast of Southwestern Johor in Malaysia

The shell height (SH) was determined by
finding the point that was the farthest along
the line perpendicular to the umbo. The shell
length (SL) was determined by measuring the
length of the perpendicular line to the umbo.
When measuring shell width (SW), the distance
from left to right was used as the reference
point (Figure 2). The estimated volume of each
pen shell (SV) was determined by multiplying
the three morphometric variables according to
Salas-Moya et al. (2014), Góngora-Gómez et al.
(2015) and García-Ulloa et al. (2019) in order
to comprehend a potential link between the
accessible spaces of pen shell for the shrimp.
Results
Snapper shrimp in the pen-shell valve were
semi-transparent and pale yellow (Figure 3).
The snapper shrimp were found and identified
from the Family Palaemoidae and genus
Anchistus and species custoides Bruce, 1977
with 15 to 20 mm length (Figure 4). The broadest
cephalothorax and the presence of eggs attached
Figure 2: Morphometric measurement of pen shell (A), Shell Height (SH), (B) Shell Length (SL) and (C)
Shell Width (SW)
Figure 3: A few examples of pen-shell habitats in the study areas. A - Pen shells (arrow) from Merambong
shoal found inhabited with seagrasses and seaweed. B - Pen shell inhabit with spoon grass Halophila ovalis

to the pleopods determined female snapper
shrimps. Usually, the width of the cephalothorax
distinguishes males and females of Anchistus
custoides [Figures 4 (B) and (D)]. The breadth of
the cephalothorax was substantially connected to
its length (rs
=0.563, p≤0.01, N=40) and females
had broader cephalothoraxes [Figure 5 (A)]. The
chelae on the second pereiopods of shrimp were
not the same size and there was a correlation
between the size of the chelae and sex [Figures
5 (B) and (C)]. The left chelae were greater than
the right chelae of female and male shrimp. The
average size of chelae of males (13.82±1.12
mm) and females (12.20 ±1.95 mm).
During this study, the symbiotic snapper
shrimp was observed inhabiting the mantle
cavity of the pen shell. Four species of pen
shell from two genera (Pinna and Atrina) have
been identified. The genus of Pinna comprising
of Pinna bicolour, Pinna muricata and Pinna
deltodes. At the same time, Atrina vexillum
represented the genus Atrina. It was found that,
out of the 40 pen shells, only 21 pen shells
were found to be inhabited by the snapper
shrimp (Table 1). A total of 40 individuals of
the snapper shrimp have been recorded. Pinna
bicolour recorded the highest number of shrimps
inhabiting the shells with 13 shrimps.
Pinna muricata with recorded the higher
SH averaged 283.94±34.5 mm while the lowest
was Pinna bicolor 216.79±53.15 mm. Pinna
muricata also recorded the highest average value
for SL141.88±11.8 mm and Pinna deltodes
recorded the lowest 105.50±27.0 mm. The
highest SW was recorded by Atrina vexillum at
44.83±6.42 mm and Pinna deltodes recorded
the lowest at 25.77±8.10 mm (Table 2). Based
on the measurements for shell volume (SV)
found, Pinna bicolour recorded a wider space
value compared to other pen shells with the
value of 2.62±1.36 dm3
. Therefore, the shrimp
that inhabits Pinna bicolour are more numerous
than the others because of the space provided.
Part of the shrimp found inhabiting the pen shell
solitary male (n=4) and solitary female (n=2).
Typically, they occur as adult pairs inhabiting
the mantle cavity.
Figure 4: Anchistus custoides Bruce, 1977 found in the mantle cavity of the pen shells. (A) and (B) are
female with (A) showing the presence of eggs attached to the pleopods and overlaying abdominal pleura
(arrow). (C) Male showing pleopods with no overlying pleura and the difference in the size of the chela of
the second periopods. Female (B) and male (D), dorsal view. Scale (5 mm)

Table 1: Distribution of snapper shrimp Anchistus custoides Bruce, 1977 in the mantle cavity of pen shells
Pen Shell
Species
Number of
Pen Shells
Inhabited by
Shrimps
Shrimp
Male Female Male and Female Total Number of Shrimps
Pinna bicolor 7 0 1 6+6 13
Pinna muricata 4 1 0 3+3 7
Pinna deltodes 5 1 0 4+4 9
Atrina vexillum 7 2 1 4+4 11
Figure 5: The proportion of breadth to the length between the cephalothorax of male and female Anchistus
custoides (A), the correlation between the length of the right chela of the second pereiopod (represented by
closed symbols) and the left chela of the second pereiopod (represented by open symbols) and the total body
length of female and male pen shell inhabitants (B) and a comparison of the length of the female and male
Anchistus custoides shells as well as their overall length (C)

Studies found that the left chela of a female
was somewhat bigger than the right chela, even
though both were smaller than that of a male of
comparable length, whose left chela was much
larger than the right. A substantial correlation
(rs
=0.450, p≤0.01, N=40) was obtained between
the total length of the pen shells and the total
length of the male and female shrimp [Figure 5
(C)]. This association suggests that the biggest
individuals occurred in the bigger pen shell
while the smallest individuals inhabited the
smaller size of the pen shell.
Discussion
A number of studies have highlighted the
relevance of the host-guest size association
between shrimps and the various types of bivalve
organisms. It was the first time that reported the
snapper shrimp Anchistus custoides resided
in the mantle cavity of pen shells in Malaysia
seagrass habitat. In the sea of Hong Kong, it
was observed that adult pairs of the pontoniine
shrimps Anchistus custos and Conchodytes
monodactylus inhabit P. bicolor (Morton, 1987).
In another situation, a shrimp, Conchodytes
nipponensis (DeHann) was found living in the
mantle cavity of a specimen of A. pectinata
from Korea (Lee & Ko, 2013) and Dutch New
Guinea (Rosewater, 1961). Garcial-Ulloa et al.
(2019) discovered that the pearl oyster shrimp
Pontonia margarita lived in the mantle cavity of
the Pinna rugosa.
This study found that the mantle cavity of
the pen shell contained both male and female
snapper shrimp. According to Richardson et
al. (1997), female snapper shrimp may be
distinguished by the presence of eggs connected
to the pleopods or in their absence, by the wider
look of the cephalothorax and the evident extent
of the overlapping abdominal pleural partly
covering the pleopods. Gracia-Ulloa et al.
(2019) reported the female cephalothorax length
was the widest compared to male cephalothorax
length with an average of 9.48±0.16 mm and
5.05±0.76 mm, respectively, being similar to the
cephalothorax length average (females = 8.21
± 2.46 mm; males = 6.39±2.02 mm) reported
by Cabrera-Pena and Solano-Lopez (1996).
A similar observation was recorded from this
study with an average of 8.80 ±0.71 mm and
6.27±0.37 mm for females and males.
Richardson et al. (1997) reported that
the chelae of the second pereiopods of P.
pinnophylax are of unequal size and there is also
a difference in size of chelae in relationship to
sex. The female’s right claw is slightly larger
than the left, although both are smaller than
those of male P. pinnophylax of similar length
Table 2: Means and ranges of shell measurements of four species of pen shells collected from
Merambong shoal
Species N
Shell Height (SH)
(mm)
Shell Length (SL)
(mm)
Shell Width (SW)
(mm)
Shell Volume
(SV)
(dm3
)
Range Mean Range Mean Range Mean Range Mean
Pinna
bicolor
15
150.11-
326.90
216.79±
53.15
65.35-
150.89
107.62±
31.77
16.12-
36.25
28.45±
8.16
0.19-
5.18
2.62±
1.36
Pinna
muricata
5
246.85-
288.84
283.94±
34.51
127.57-
153.59
141.88±
11.80
25.63-
33.25
29.15±
4.18
0.95-
1.41
1.16±
0.20
Pinna
deltodes
10
115.57-
276.81
203.44±
48.10
61.62-
127.90
105.50±
20.27
15.36-
32.12
25.77±
8.10
0.25-
1.17
0.62±
0.36
Atrina
vexillum
10
164.10-
224.78
192.65±
22.74
107.83-
155.61
131.07±
16.55
38.16-
61.09
44.83±
6.42
0.74-
1.52
1.18±
0.43
Note: N = number of samples

and whose right chela is substantially larger
than the left. Richardson et al. (1997) found
that the chelae of the second pereiopods of P.
pinnophylax are not similar and there is also
a size disparity between the chelae and the
gander. Although the right claw of the female
P. pinnophylax is marginally bigger than the
left, both claws are much shorter than those
of male P. pinnophylax of comparable length,
whose right chela is significantly larger than the
left. The presence study found the left chelae
of Anchistus custoides was greater than right
chelae and the average length of male chelae
was greater than female. According to García-
Ulloa et al. (2019), the maximum chelae length
of the second pereiopod of P. margarita females
was longer than their male counterparts with an
average of 15.61±1.94 mm and 12.65±3.03 mm.
During the study conducted on the
Merambong shoal, pen shell was found living
in muddy sand areas and associated with
seagrasses. Seagrass areas provide an important
source of food for aquatic life. In the Sungai
Pulai estuary, (Hossain et al., 2018) reported
ten species of seagrasses inhabiting the sandy-
muddy area while Arina et al. (2020) recorded
seven species (Cymodocea serrulata, Halophila
ovalis, Halodule pinifolia, Enhalus acoroides,
Thalassia hemprichii, Halodule uninervis
and Syringodium isoetifolium) of seagrasses
inhabiting the sandy-mud area in Merambong
shoal. According to Idris et al. (2009), a total of
seven species of pen shell have been recorded at
Merambong shoal but from this study, only four
species of pen shell have been recorded.
From the study conducted, it was found
that the presence of Anchistus custoides
inhabiting four species of pen shells. Pinna
bicolour recorded higher space than three other
species with a range size of 150.11-326.90 mm
and an average of 216.79±53.15 mm with the
SV 2.62±1.36 dm3
. García-Ulloa et al. (2019)
reported the presence of shrimp in pen shells
with SH ranging from 198 to 271 mm. However,
there was no association between the size of the
shrimp and the host shell. This research found
that the average estimated volume for each
rugose pen shell (1.26±0.76 dm3
) was lower than
what was reported for A. tuberculosa (3.55±0.76
dm3
) by Góngora-Gómez et al. (2015) using the
same morphometric computation. This would
imply that the SV of the Pinna bicolour was
big enough to accommodate not only a single
Anchistus custoides but also a male and female.
However, Baeza et al. (2013) conclude that a
shell length of <175 mm is insufficient for the
symbiotic shrimp. Aucoin and Himmelman
(2010) came to a similar conclusion when they
investigated the development of Pontonia sp.
with its host, the pin shell Pinna carnea.
The investigations of the shrimps indicated
that they react either violently or defensively
along the shell edge, and they might serve as a
warning to any curious predatory fish prevalent
within the canopy of the seagrass meadow
(Bell & Harmelin-Vivien, 1982; 1983). On the
other hand, it is unclear whether the Anchistus
custoides benefit from the association. In a
study of the zoea development of Pontonia
pinnophylax. Calafiore et al. (1991) found that
the development from zoea stage VIII to the
post-larval stage only occurred in the presence of
adult mussels. In the absence of Pinna, the zoea
continued to grow but they did not transform
into juvenile shrimps.
According to Richardson et al. (1999),
the snapper shrimp, pontoniine demonstrated a
strong affinity for shade and they rapidly became
immobilised in even a weak current flow.
Therefore, the presence of these shrimp inside
pinnids would provide them with the necessary
shelter, shade and protection. The shrimp may
get some of their nutrients from the pen shells.
When Pinna is being fed in suspension, there is
never a break in the creation of pseudofaeces
on its part. In most cases, the contractions of
the adductor muscle are responsible for the
expulsion of pseudofeces; however, it seems
that this is not the case in Pinna (Yonge, 1953).
It is currently unclear whether or if shrimp are
there, what function they play or what kind
of influence they have on the pen shell. The
presence of shrimp at the shell border and
patrolling the mantle margin may operate as a

first line of defense, inhibiting predatory fish
from grazing on the posterior edge of the mantle
and shell (Richardson et al., 1999). The precise
nature of the interaction between the two species
has not been scientifically established in this
research. This is because Anchistus custoides
represents the last link in a series of species that
are all at risk of extinction. In the coastal seas
of Johor, Malaysia, there is a kind of shrimp
that lives inside of a pen shell, which is also an
endangered bivalve.
Conclusion
A pair and single of adult snapper shrimp
Anchistus custoides Bruce, 1977 with the ranges
size of 15 to 20 mm in length have been recorded
inhabiting the mantle cavity of pen shell in
Merambong shoal seagrass beds. Pinna bicolour
and Atrina vexillum recorded the highest number
of shrimp inhabiting the shell with the average
SH 150.11-326.90 mm with the SV 2.62±1.36
dm3
and SH 164.10-224.78 mm with the SV
1.18±0.43 dm3
, respectively. It was observed
that the shrimp was living symbiotically with
the pen shells. Female shrimps were identified
by the presence of eggs attached to the pleopods
and the female size was greater than the male
specimens.
Adult snapper shrimp Anchistus custoides
has been found living in the mantle cavity of
pen shells in the seagrass beds of Merambong
shoal. Their lengths vary from 15 to 20 mm. A
pair and a single adult snapper shrimp Anchistus
custoides Bruce, 1977 have been reported.
Pinna bicolour and Atrina vexillum reported
the maximum number of shrimp occupying the
shell, with an average shell height of 150.11-
326.90 mm and a shell volume of 2.621.36
dm3
for Pinna bicolour and SH 164.10-224.78
mm and SV 1.18±0.43 dm3
for Atrina vexillum,
respectively. It was also discovered that the
shrimp lived symbiotically with the pen shells.
It was possible to determine whether or not a
shrimp was female by seeing whether or not it
had eggs attached to its pleopods. Additionally,
the size of female specimens was larger than that
of male specimens.
Acknowledgements
The author would like to acknowledge the
Ministry of Higher Education Malaysia (FRGS)
research grant 2014. The author also would
like to thank the deanery and staff from the
Department of Animal Science and Fishery,
Faculty of Agricultural and Forestry Sciences,
Universiti Putra Malaysia Bintulu Sarawak
Campus for technical, logistic and facilities
support which made this study possible.
References
Anker, A. & De Grave, S. (2016). An update and
annotated checklist of marine and brackish
caridean shrimps of Singapore (Crustacea,
Decapoda). Raffles Bulletin of Zoology, 30,
343-454.
Arina, N., Ashikin, C. N., Hidayah, N., Fairoz,
M., & Rozaimi, M. (2020). Data on
sediment nitrogen loadings and nitrogen
in the biomass of seagrasses from Sungai
Pulai estuary (Johor, Malaysia). Data in
Brief, 28, 104979. https://doi.org/10.1016/j.
dib.2019.104979
Aucoin. S. & Himmelman, J. H. (2010). A first
report on the shrimp Pontonia sp. and other
potential symbiont in the mental cavity of
the pen shell Pinna carnea in the Dominica
Republic. Symbiosis, 50, 135-141.
Baeza, J. A. (2008). Social monogamy in the
shrimp Pontonia margarita, a symbiont of
Pinctada mazatlanica, off the Pacific coast
of Panama. Marine Biology, 153, 387-395.
Baeza, J. A. (2010). The symbiotic lifestyle
and its evolutionary consequences: Social
monogamy and sex allocation in the
hermaphrodite shrimp Lysmata pederseni.
Naturewissen-schaften, 97(8), 729-741.
Baeza, J. A., Bolanos, J. A., Hernandez, J. E.
Lira, C. & Lopez, R. (2011). Monogamy
do not last long in Pontonia mexicana,
a symbiotic shrimp of the Amber pen
shell Pinna carnea from the Southeastern
Caribbean Sea. Journal of Experimental
Marine Ecology, 407(1), 41-47.

Baeza, J. A., Ritson-William, R. & Fuentes, M.
S. (2013). Sexual and mating system in the
caridean shrimp symbiotic in the winged
pearl oyster in the Coral Triangle. Journal
of Zoology. 289, 172-181.
Baeza, J.A., Hemphill, C.A., & Ritson-
Williams, R. (2015). The sexual
and mating system Odontonia katoi
(Palemonidae, Pontoniinea), a symbiotic
guest of the ascidian Polycarpa aurata
in the Coral Triangle. PLOS ONE, 10(3),
e0121120. https://doi.org/10.1371/journal.
pone.0121120
Bell, J. D & Harmelin-Vivien, M. L. (1982). Fish
fauna of France Mediterranean Posidonia
oceanica seagrass meadows. 1. Community
structure. Téthys. 10, 1-14.
Bell, J. D & Harmelin-Vivien, M. L. (1983). Fish
fauna of France Mediterranean Posidonia
oceanica seagrass meadows. 2. Feeding
habits. Téthys. 11, 337-347.
Bhujel, R. C. (2008). Statistics for aquaculture.
Ames, Wiley-Blackwell, 222 p.
Butler, A., N. Vicente & De Gaulejac, B.,
(1993). Ecology of the Pterioid Bivalves
Pinna bicolor Gmelin and Pinna nobilis L.
Mar. Life. 3(1-2), 37 - 45.
Butler, A. J. (1987). Ecology of Pinna bicolor
Gmelin (Mollusca: Bivalvia) in Gulf
St Vincent, South Australia: Density,
reproductive cycle, recruitment, growth and
mortality at three sites. Australian Journal
of Marine and Freshwater Research. 38,
743-769.
Butler, A. J. & Brewster, F. J. (1979). Size
Distribution and growth of the fan-
shell Pinna bicolor Gmelin (Mollusca:
Eulamellibranchia) in South Australia.
Australian Journal of Marine and
Freshwater Research, 30(1), 25-39.
Butler, A. J., & Keough, M. J. (1981).
Distribution of Pinna bicolor Gmelin
(Mollusca: Bivalvia) in SouthAustralia with
observations on recruitment. Transactions
of the Royal Society of South Australia,
Incorporated, 105(1): 29-39.
Cabrera-Pena, J., & Solano-Lopez, Y. (1996).
Growth of Modiolus capax (Bivalvia:
Mytilidae) by means of suspended culture
system. Revista de Biología Tropical, 44,
939-940.
Calafiore, N., Costancxzo, G. & Giacobbe, S.
(1991). Mediterranean species of the genus
Pontonia Latreille, 8129. 1. Developmental
stages of Pontonia pinnophylax (Otto,
1821) (Decapoda, Natantia, Pontoniinae)
reared in the laboratory. Crustaceana, 60,
52-75.
Chow, L. H., De Grave, S., & Tsang, L. M.
(2021). Evolution of protective symbiosis
in palaemonid shrimps (Decapoda:
Caridea) with emphases on host spectrum
and morphological adaptations. Molecular
Phylogenetics and Evolution, 162,
107201. https://doi.org/10.1016/j.ympev.
2021.107201
Cummings, V. J., Thrush, S. F., Hewitt, J. E. &
Turner, S. J. (1998). The influence of the
pinnid bivalve Atrina zelandica (Gray) on
benthic macroinvertebrate communities
in soft sediment habitats. Journal of
Experimental Marine Biology and Ecology,
228, 227-240.
De Grave S, & Fransen C.H.J.M. (2011).
Carideorum catalogus: The recent species
of the dendrobranchiate, stenopodidean,
procarididean and caridean shrimps
(Crustacea: Decapoda). Zoologische
Mededelingen Leiden, 85(9), 195-589.
Dobson, N. C., De Grave, S., & Johnson,
M. L. (2014). Linking eye design
with host symbiont relationships in
pontoniine shrimps (crustacea, decapoda,
palaemonidae). PLOS ONE, 9(6), e99505.
https://doi.org/10.1371/journal.pone.0099
505
Garcia-Ulloa, D., Landa-Jaime, V., Gongora-
Gomez, A. M., Garcia-Ulloa, M., &
Hernandez-Sepulveda, J. A. (2019). Sexual

and reproductive traits of the oyster
shrimp Pontonia margarita (Decapoda:
Palemonidae), Symbiotically inhabiting the
mantal cavity of the rugose pen shell Pinna
rugosa (Bivalvia: Pinnidae). Zoologia, 36,
e29774.
Gongora-Gomez, A. M., Munoz-Sevilla, N.
P., Hernandez-Sepulveda, J. A., Garcia-
Ulloa, II, M & Garcia-Ulloa, M. (2015).
Association between pen shell Atrina
tuberculosa and the shrimp Pontonia
margarita. Symbiosis, 66(2), 107-110.
Hossain, M. S., Hashim, M., Bujang, J. S.,
Zakaria, M. H., & Muslim, A. M. (2018).
Assessment of the impact of coastal
reclamation activities on seagrass meadows
in Sungai Pulai estuary, Malaysia, using
Landsat data (1994-2017). International
Journal of Remote Sensing, 40(9), 3571-
3605. https://doi.org/10.1080/01431161.20
18.1547931
Idris, M. H., Arshad, A., Amin, S. M. N. N.,
Japar, S. B., Daud, S. K., Mazlan, A. G.,
Zakaria, M. S., & Yusoff, F. M. (2012).Age,
growth and length-weight relationships of
Pinna bicolor Gmelin (Bivalvia: Pinnidae)
in the seagrass beds of Sungai Pulai Estuary,
Johor, Peninsular Malaysia. Journal of
Applied Ichthyology, 28(4), 597-600.
https://doi.org/10.1111/j.1439-0426.2011.
01807.x
Idris, M. H., Arshad, A., Japar Sidik, B.,
Mazlan, A. G., & Daud, S, K. (2008). New
distribution of two pen shells (Bivalvia:
Pinnidae) from the seagrass beds of
Sungai Pulai, Johor, Malaysia. Journal of
Biological Sciences, 8(4), 120-132.
Idris, M. H.,Arshad,A., Japar Sidik, B., Mazlan,
A. G., & Daud, S, K. (2009). Morphological
characteristics of Pinna bicolor Gmelin and
Pinna deltodes Menke from the seagrass
bed of Sungai Pulai, Johore, Peninsular
Malaysia. Sains Malaysiana, 38(3), 333-
339.
Itani, G., Kato, M. & Shiriyama, Y. (2002).
Behaviour of the shrimp ectosymbionts,
Peregrinamor ohshimai (Mollusca:
Bivalvia) and Phyllodurus sp. (Crestacean:
Isopoda) through host ecdysis. Journal
of the Marine Biology Association of the
United Kingdom, 82, 69-78.
Katsanevakis, S. (2004). Population ecology of
the endangered fan mussel Pinna nobilis
in the marine lake. Endangered Species
Research. 1, 1-9.
Kennedy, H., Richardson, C.A., Duarte, C.M., &
Kennedy, D.P. (2001). Diet and association
of Pontonia pinnophylax occurring in
Pinna nobilis: Insights from stable isotope
analysis. Journal of the Marine Biology
Association of the United Kingdom, 81,
177-178.
Keough, M. J. (1984a). Effect of patch size on the
abundance of sessile marine invertebrate.
Ecology. 65(2), 423-437.
Keough, M. J. (1984b). Dynamics of the
Epifauna of the Bivalve Pinna bicolor:
Interactions among Recruitment, Predation,
and Competition. Ecology, 65(3), 677-688.
Lee, K. H., & Ko, H. S. (2011). First records
of two Pontoniid shrimps (Crustacea:
Decapoda: Caridea: Palaemonidae) from
Korea. Animal Systematics, Evolution
and Diversity, 27(3), 239-245. https://doi.
org/10.5635/kjsz.2011.27.3.239
Lee, S.K. & Ko, H.S. (2013). The first record of
Conchodytes nipponensis (De Haan, 1844)
(Crustacea: Decapoda: Palaemonidae)
associated with pen shell Atrina pectinata
(Linnaeus, 1767) (Mollusca: Bivalvia) from
Korea. Journal of Species Research, 2(1),
1-6.
Morton, B., (1987). Temporal host segregation
by Anchistus custos and Conchodytes
monodactylus (Crustacea: Pontoniinae) of
Pinna bicolor (Bivalvia: Pinnidae) in Hong
Kong. Asian Marine Biology. 4, 129-140.
Munguia, P. (2004). Successional patterns on
pen shell communities at local and regional
scales. British Ecological Society. Journal
of Animal Ecology. 73, 64-74.

Nizinski, M. S. (1989). Ecological, distribution,
demography, and behavioural observations
on Periclimens anthophilus, an atypical
symbiotic cleaner shrimp. Bulletin of
Marine Sciences, 45, 174-188.
Olliff, E. R. R. (2013). Symbiosis of the sea star
shrimp, Periclimenes soror Nobili, 1904
(Decapoda, Palaemonidae), and cushion
star, Culcita novaeguineae Müller &
Troschel, 1842 (Echinodermata,Asteroidea,
Oreasteridae): Host finding and benefits.
Crustaceana, 86(5), 564-577. https://doi.
org/10.1163/15685403-00003192
Overstreet, R. M., & Lotz, J. M. (2016). Host-
symbiont relationships: Understanding
the change from guest to pest. In C. J.
Hurst (Ed.), The Rasputin effect: When
commensals and symbionts become
parasitic (Advances i, pp. 27-64). Springer
International Publishing. https://doi.
org/10.1007/978-3-319-28170-4_2
Piero, D. F & Mantelatto, F.L. (2011). Population
dynamic of the pea crab Austinixa aidae
(Brachyura, Pinnotheridae): A symbiotic
of the ghost shrimp Callichirus major
(Thalassinidae, Callianassidae) from the
southwestern Atlantic. Iheringia, 101, 5-14.
Poutiers, M. (1998). The living marine resources
of the Western central Pacific. In K.
Carpenter & V. Niem (Eds.), FAO species
identification guide for fishery purposes.
The living marine resources of the Western
Central Pacific. Volume 2. Cephalopods,
crustaceans, holothurians and sharks.
[classification of PORTUNIDAE ] FAO
Species Identification Guide for Fishery
Purposes (1st
ed.). Food and Agriculture
Organization of the United Nations.
Radda, B., & Milat, T. (2009). The first record of
thedecapod,Pontoniapinnophylax,inPinna
nobilis from the southern Adriatic (Croatia)
(Decapoda, Natantia). Crustaceana, 82(11),
1383-1392. https://doi.org/10.1163/001121
609X12475745628469
Rabaoui, L., Zouari, S.T., & Hassine, O.K.B.
(2008). Two species of Crustacea
(Decapoda) associated with the fan mussel,
Pinna nobilis Linnaeus, 1758 (Mollusca:
Bivalvia). Crustaceana. 81:433-446.
Richardson, C. A., Kennedy, H., Duarte, C.M.,
Kennedy, D.P. & Proud. S. V. (1997).
The occurrence of Pontonia pinnophylax
(Decapoda: Natantia: Pontoniinae) in Pinna
nobilis (Mollusca: Bivalvia: Pinnidae) from
the Mediterranean. Journal of the Marine
Biological Association of the United
Kingdom, 77, 1227-1230.
Richardson, C. A., Kennedy, H., Duarte, C. M.,
Kennedy, D. P. & Proud. S. V. (1999). Age
and growth of the fan mussel Pinna nobilis
from south-east Spanish Mediterranean
seagrass (Posidonia oceanica) meadows.
Marine Biology, 133, 205-212.
Rosewater, J. (1961). The Family Pinnidae in
the Indo-Pacific. Indo-Pacific Mollusca.
1(4), 175-226.
Rosewater, J. (1982). Review of Hawaiian
Pinnidae and revalidation of Pinna exquisite
Dall, Bartsch, and Rehder, 1938 (Bivalvia:
Mytiloida). Pacific Science. 36(4), 453-458.
Salas-Moya, C., Mena, S. & Wehrtmann,
I. S. (2014). Reproductive traits of the
symbiotic pea crab Austinotheres angelicus
(Crustacea: Pinnotheridae) living in
Saccostrea palmula (Bivalvia: Ostreidae),
Pacific coast of Costa Rica, Zookeys, 457,
239-252.
Scheltema, A. (1983). Pinna deltodes Menke
newly described and differentiated from
P. bicolor Gmelin (Bivalvia, Pterioida).
Journal of the Malacological Society of
Australia, 6(1-2), 37-52. https://doi.org/10.
1080/00852988.1983.10673953
Silentic, T. & Peharda, M. (2003). Population
study of the fan shell Pinna nobilis L.
In Malo and Veliko Jezero of the Mljet
National Park (Adriatic Sea). Scientia
Maria, 67(1), 91-98.
Yonge, C.M. (1953). Form and Habit in Pinna
carnea Gmelin. Philosophical Transactions
of the Royal Society of London, Series B.
237(648), 335-374.

Zavodnik, D. (1967). Contribution to the ecology
of Pinna nobilis L. (Mollusca: Bivalvia) in
the northern Adriatic. Thalassia Jugosl, 3,
93-102.
Zavodnik, D., Hrs-Brenko, M. & Legac, M.
(1991). Synopsis on the fan shell Pinna
nobilis L. in the eastern Adriatic Sea. In
C.F. Boudouresque, M. Avon and V. Gravez
(Eds.), Les Especes Marines a Proteger
en Mediterranee. GIS Posidonie Publ.,
Marseille, France. 169-178.

Symbiotically snapper shrimp Anchistus custoides Bruce, 1977 (Decapoda: Palaemonidae) inhabiting the mantle cavity of the pen shell (Bivalvia: Pinnidae)

Recommended

Recommended

More Related Content

Similar to Symbiotically snapper shrimp Anchistus custoides Bruce, 1977 (Decapoda: Palaemonidae) inhabiting the mantle cavity of the pen shell (Bivalvia: Pinnidae)

Similar to Symbiotically snapper shrimp Anchistus custoides Bruce, 1977 (Decapoda: Palaemonidae) inhabiting the mantle cavity of the pen shell (Bivalvia: Pinnidae) (20)

More from AbdullaAlAsif1

More from AbdullaAlAsif1 (20)

Recently uploaded

Recently uploaded (20)

Symbiotically snapper shrimp Anchistus custoides Bruce, 1977 (Decapoda: Palaemonidae) inhabiting the mantle cavity of the pen shell (Bivalvia: Pinnidae)