688 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 688-701, e46811, enero-diciembre 2022 (Publicado Set. 30, 2022)

Seagrass density correlates with burrow abundance and size

in the Zebra Mantis Shrimp (Stomatopoda: Lysiosquillidae)

Nadiarti Nurdin Kadir1*; https://orcid.org/0000-0003-2837-1914

Yayu Anugrah La Nafie2; https://orcid.org/0000-0002-7461-0034

Dody Priosambodo3; https://orcid.org/0000-0002-1817-9243

1. Aquatic Resource Management Study Program, Fisheries Department, Universitas Hasanuddin, Makassar, Indonesia;

nadiarti@unhas.ac.id (*Correspondence)

2. Marine Science Study Program, Marine Science Department, Universitas Hasanuddin, Makassar, Indonesia;

yayulanafie@yahoo.com

3. Biology Study Program, Biology Department, Universitas Hasanuddin, Makassar, Indonesia;

d.priosambodo@unhas.ac.id

Received 21-IX-2021. Corrected 09-III-2022. Accepted 21-IX-2022.

ABSTRACT

Introduction: Mantis shrimps are ecologically and economically important organisms in marine ecosystems.

However, there is still a lack of information about their habitat, in particular, their burrows.

Objective: To analyze how dense and sparse mantis shrimp burrows differ in abundance, size, sediment grain

size, and water quality.

Methods: We counted burrows in 10 x 10 m2 random plots in sparse and dense seagrass (ten plots per density),

around Barrang Lompo Island, South Sulawesi, Indonesia. Sampling took place at spring low tide from August

to September 2017.

Results: Two mantis shrimp species were observed: Lysiosquillina maculate and L. sulcata. Dense and sparse

seagrass burrows did not differ in wall grain size or water parameters, both inside and outside of the burrows

(P > 0.05). Similarly, there was no correlation between burrow depth and diameter in either dense (P > 0.05; r=

0.27) or sparse (P > 0.05; r= 0.33) seagrass. However, larger burrows tend to occur in denser beds, but there were

more burrows in denser seagrass (t-test, P < 0.05).

Conclusions: There seems to be a preference for dense seagrass beds, especially by larger mantis shrimps. The

correlation between shrimp burrow abundance and seagrass density highlights the importance of conserving the

quality as well as the extent of seagrass habitat.

Key words: Zebra mantis shrimp; burrows; seagrass; intertidal; Barrang Lompo Island; Indonesia.

https://doi.org/10.15517/rev.biol.trop.2022.46811

AQUATIC ECOLOGY

INTRODUCTION

Mantis shrimps (Phylum Crustacea, Order

Stomatopoda) are common and economically

important coastal fisheries targets in Europe,

and even more so in Asia (Ragonese et al.,

2012; Wortham, 2009). They are used for

consumption, animal feed, and also for envi-

ronmental biomonitoring (Wortham, 2009).

Despite their economic importance, there is

very little information on the habitat fac-

tors that determine their abundance, distribu-

tion and size. In tropical Asia, these shrimps

are principally harvested in shallow seagrass

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beds and other soft-sediment habitats (Mili

et al., 2013; Taylor & Patek, 2010), in which

they inhabit burrows. Hence, understanding the

importance of these burrows for mantis shrimp

abundance, distribution and size will be valu-

able for managing sustainable harvesting.

Mantis shrimps are known as predatory

burrowing crustaceans. These animals can be

found over a wide depth range, from shallow

and intertidal waters to subtidal areas tens of

meters deep (Poupin & Poupin, 2008). Like

other burrowing animals, their burrows provide

protection from predators and space for repro-

duction (Mead & Minshall, 2012; Morgan &

Goy, 1987). The burrows also serve an aeration

function, preventing anaerobic conditions by

increasing the area of the water-sediment inter-

face and ventilating the lower sediment layers

with oxygen-rich water (Kinoshita, 2002). A

limited number of studies have been carried out

on mantis shrimp burrows. For example, chem-

ical communication related to cavity occupa-

tion in Gonodactylus festai (Caldwell, 1979),

burrow opening size of the mantis shrimp

Squilla empusa (Mead & Minshall, 2012),

occupation of natural and artificial burrows by

Japanese mantis shrimp (Oratosquilla orato-

ria) (Matsuura & Hamano, 1984), aggressive

territorial defense behavior of Gonodactylus

bredini (Dingle & Caldwell, 1969). But none

of these studies focus on mantis shrimp abun-

dance in seagrass beds or the factors influenc-

ing their abundance.

Seagrass beds are well known as an impor-

tant habitat harboring a variety of fauna (Nadi-

arti et al., 2015), providing food, protection,

shelter, and living space (Christianen et al.,

2013; Jackson et al., 2001; Jackson et al., 2006;

Vonk et al., 2010). The seagrasses located in

the intertidal zone are more frequently exposed

to anthropogenic stress (Benjamin et al., 2008;

Crowe et al., 2000) and seagrass in Indonesia

have experienced widespread degradation and

loss (Nadiarti et al., 2012; Unsworth et al.,

2018). Associated organisms in this ecosystem,

including mantis shrimps, are therefore coming

under pressure and potentially threatened.

Zebra mantis shrimp can be found in a

variety of habitat types, including soft substrate

such as mud (Reaka, 1987) bare sandy bottoms,

seagrass beds and coral rubble-dominated areas

(Priosambodo et al., 2014), as well as in coral

reef ecosystems (Barber et al., 2011). However,

information is still extremely limited on their

dwellings (burrows) in general, in seagrass

beds in particular, and specifically with respect

to seagrass density. Therefore, in this study we

ask the following questions: i) does the density

of seagrass beds influence burrow abundance;

ii) does the sediment grain size affect burrow

wall construction in seagrass beds of differ-

ent density; iii) does water quality (dissolved

oxygen, pH and temperature) differ between

the outside and inside of mantis shrimp bur-

rows in seagrass beds of different density. To

answer these questions, we analyzed the differ-

ences between dense and sparse seagrass beds

in burrow abundance, burrow size distribution

and correlation between diameter and depth of

the burrows, as well as the differences in water

quality (dissolved oxygen, pH and temperature)

outside and inside mantis shrimp burrows.

MATERIALS AND METHODS

Field sampling and data collection:

Mantis shrimp burrows were identified during

field observation when the mantis shrimp was

visible in the burrow and following previously

described burrow characteristics (Ramsay &

Holt, 2001). Shrimp burrows were studied at

two seagrass sites within the intertidal waters

around Barrang Lompo Island (5º03’ S-119º19’

E, 1.5 m.s.n.m) in the Spermonde Archipelago,

South Sulawesi, Indonesia (Fig. 1). Sampling

took place at spring low tide from August to

September 2017, i.e. during the calm season at

this site, to ensure good visibility and accessi-

bility for taking measurements of both the bur-

row dimensions and water quality parameters.

At site 1 on the West coast (50 x 300 m),

the seagrass beds were classified as sparse (20-

50 % cover), while at site 2 on the Southwest

coast (50 x 300 m) the seagrass beds were

classified as dense (> 80 % cover). The species

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present were also identified following McK-

enzie (2003). The seagrass percentage cover

at each site was estimated using a standard

percentage cover photography-based method

(Mckenzie, 2003).

Shrimp species identification is important

for controlling and managing the exploitation

of mantis shrimps. Mantis shrimps were not

extracted from their burrows or caught; how-

ever, all mantis shrimps seen in the study area

(inside or outside the burrows) were identified

based on the descriptions and keys in Ahyong

(2001), Ahyong and Randall (2001) and Man-

ning (1978). Particular features of interest for

identification were the number of teeth on the

dactylus of the raptorial claw and the coloration

of the distal end of the uropodal endopod.

Ten plots (10 m x 10 m each) were placed

randomly in each of the two sites. Distance

between plots was determined based on the

number of steps obtained from a Random

Integer Generator (RIG). In each plot, the

mantis shrimp burrows were counted and their

diameter and depth recorded with a folding

plastic ruler which was narrow enough to avoid

touching or disturbing the sides of the burrow.

A yabbie pump was not used in order to avoid

undue disturbance to the burrow or the shrimp.

Some mantis shrimps have U-shaped burrows

(Matsuura & Hamano, 1984), although the dia-

gram in Faulkes (2013) indicates a straight bur-

row for the genus Lysiosquilla. The ruler was

inserted gently into the burrow until the bottom

or (if present) the U bend of the burrow was

encountered. In order to compare characteris-

tics inside and outside burrows, we measured

sediment grain size composition and water

quality parameters (temperature, pH, and dis-

solved oxygen) inside and outside each burrow.

Sediment samples (to a depth of ca. 15-20

cm) from outside the burrows were collected

using a homemade yabbie pump; and sedi-

ment samples from inside the burrows (wall

lining) were taken to depths of between 6-15

cm (depending on burrow depth) using a

homemade small tube with a fine screen at the

bottom of the tube. All visible animal and plant

fragments were removed from the sediment

Fig. 1. Location of the sparse (site 1) and dense (site 2) seagrass study sites around Barrang Lompo Island, Spermonde

Archipelago, South Sulawesi, Indonesia. The map of the Spermonde Archipelago was adapted from Stapel (2001).

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samples prior to aeration. Classification of

grain size based on Wen-worth Class and grain

size composition of sediment (outside burrows)

and burrow lining material (inside burrows)

was analyzed after shaking the dried sediment

sample for 10 minutes on a series of test sieves

(Din 4188 Pruf-sieb) with mesh sizes of 2, 1,

0.5, 0.25, 0.1, and 0.063 mm.

Dissolved oxygen, pH, water tempera-

ture outside and inside burrows were mea-

sured in-situ using a CellOx 325 sensor head

and a SenTix 21 sensor head, respectively.

Both sensor heads were connected to a WTW

Multi 340i Multimeter.

Data analysis: The number of burrows

per plot gave burrow abundance per 100 m2.

Statistical analyses were conducted in SPSS 25

(IBM Corp., 2017), and the summary statistics

presented as mean ± standard error (SE). The

significance of differences in mean value and

variance of burrow abundance between dense

and sparse seagrass beds was determined using

a Student’s t-test.

Non-parametric Spearman correlation

analysis was used to evaluate the significance

of correlation between burrow diameter and

depth. Burrows size distributions in the dense

and sparse seagrass beds were compared using

a Student’s t-test. Sediment grain size compo-

sition both outside and inside burrows, in dense

and sparse seagrass beds, were compared using a

factorial MANOVA analysis.

The measured water parameters were

pooled together for outside and inside burrows

from both seagrass beds. Water quality param-

eters (DO concentration, pH, and water tem-

perature) outside and inside burrows in sparse

and dense seagrass beds were analyzed using a

factorial MANOVA analysis.

RESULTS

Out of the 16 seagrass species reported

from Indonesia (Yasir & Moore, 2021), six were

observed at the study sites. Four species were

present at both sites: Cymodocea rotundata,

Enhalus acoroides, Syringodium isoetifolium,

and Thalassia hemprichii. Halophila ovalis

was found only in the sparse seagrass beds

(Site 1) and Halodule uninervis in the dense

seagrass beds (Site 2). At the sparse site, the

most visually prominent feature was the high

canopy formed by the large leaves of E. acoroi-

des. At the dense site isolated small stands of E.

acoroides were present but the general canopy

height was much lower, with Thalassia hemp-

richii the visually dominant species.

A total of 77 mantis shrimp burrows were

identified during the study. The number of

burrows can be considered a good indication

of relative abundance, as most of the burrows

found during the study were occupied by man-

tis shrimps. Zebra mantis shrimps were often

visible in their burrow openings and sometimes

came out from their burrows, enabling identi-

fication as members of the genus Lysiosquil-

lina. However, only six individuals (mostly

from larger burrows 4-6 cm in diameter) could

be observed in sufficient detail to determine

the species, comprising five Lysiosquillina

maculata and one L. sulcata. The abundance of

mantis shrimp burrows in sparse and dense sea-

grass beds was significantly different (t = 2.3,

Fig. 2. Mean number of mantis shrimp burrows in 10 x 10

m plots (100 m2) with different seagrass densities (Error

bars represent standard error).

d.f. = 18, P < 0.05), being around twice as high

in the dense seagrass beds as in the sparse sea-

grass beds (Fig. 2).

There was a significant difference between

the sparse and dense seagrass beds in both

burrow diameter (t= 5, d.f.= 76, P < 0.0001)

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and burrow depth (t= 2.7, d.f.= 76, P < 0.05).

Burrows in the sparse seagrass beds were,

on average, smaller compared to those in the

dense beds (Fig. 3). Burrow diameter and depth

ranges were 1.3-2.5 cm and 7.0-19.0 cm in the

sparse seagrass beds and 2.5-6.0 cm and 5.0-

26.0 cm in the dense seagrass beds. However,

there was no significant correlation (P > 0.05)

between burrow diameter and depth within

either the dense or sparse seagrass beds, or

indeed for the combined burrow dataset.

Overall, the sediment comprised sand,

coral rubble, and shell fragments; however, we

did not analyze the proportion of each compo-

nent. Factorial MANOVA analysis showed that

the grain size composition differed significant-

ly between the sparse and dense seagrass beds

as well as between the burrow lining materials

and the sediment outside the burrows in both

seagrass beds (Fig. 4). Inside the burrows, the

overall pattern was similar in both seagrass

beds, although there were significant differ-

ences between dense and sparse seagrass beds

in the proportions of three grain size classes:

> 2 mm, < 0.13 mm, and < 0.063 mm. The

proportion of coarse particles (> 2 mm) in the

burrow lining was significantly higher in the

sparse seagrass bed while the proportions of

fine particle classes (< 0.13 mm and > 0.063

mm) were higher in the dense seagrass bed.

Outside the burrows, the grain size distribu-

tion was noticeably different between the two

seagrass beds, and the sediment grain size

composition patterns were reversed between

the dense and sparse seagrass beds. There was

a significantly higher proportion of relatively

coarse grain classes (> 2 mm and > 0.5 mm)

in the sparse seagrass bed, while the proportion

of smaller grain size classes (> 0.25 mm and >

0.13 mm) was significantly higher in the dense

seagrass bed.

TABLE 1

Water parameters outside and inside burrows in dense and sparse seagrass beds. Values are presented as mean ± SE (n)

Seagrass beds Burrows O2 (mg l-1) pH Temperature (°C)

Dense Outside 4.51 ± 0.51 (10) 7.66 ± 0.03 (9) 30.16 ± 0.14 (10)

Inside 4.03 ± 0.58 (10) 7.59 ± 0.03 (9) 30.25 ± 0.10 (10)

Sparse Outside 5.37 ± 0.56 (7) 7.79 ± 0.08 (7) 30.28 ± 0.12 (8)

Inside 3.79 ± 0.77 (7) 7.77 ± 0.04 (7) 30.29 ± 0.09 (8)

Fig. 3. Distribution of burrows based on mean diameter (cm) and mean depth (cm). Error bars represent standard error.

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The water quality parameters measured

(DO concentration, pH, temperature) (Table 1)

did not differ significantly (factorial MANO-

VA) between the water column (outside bur-

rows) and in the burrow water (inside burrows)

or between the two seagrass bed types (sparse

and dense).

DISCUSSION

The two species observed at the study

site are spearers. In contrast with the smasher

mantis shrimp that live in pre-existing cavities

within hard substrate, the spearers live in bur-

rows that they excavate themselves, typically in

soft substrates (Hernáez et al., 2011). Spearer

mantis shrimp are ambush predators with elon-

gated spear-like appendages used to ambush

soft-bodied evasive prey (deVries et al., 2012;

deVries, 2017). Spearing appendages are more

commonly found in lysiosquillids, but also in

the squillid, bathysquillid, and some gonodac-

tylid mantis shrimps (Caldwell and Dingle,

1975). While Lysiosquillina maculata is a

widespread and frequently abundant across the

Indo-Pacific (Ahyong, 2001), including some

areas of Indonesia (e.g. Dini et al., 2013), L.

sulcata is only reported from a few locations;

however, these range from the Pacific to Mada-

gascar and Zanzibar including one Indonesian

site in Ambon (Manning, 1978); French Poly-

nesia, including Moorea (Patek et al., 2012) and

Rangiroa (Lecchini et al., 2010); and Australia,

including One Tree Island, Australia (Courtney

et al., 1999; Courtney et al., 2007). While the

literature on L. sulcata is extremely limited, it

has been reported as sympatric with L. macu-

lata (Manning, 1978; Lecchini et al., 2010).

Like other spearer mantis shrimps, the species

L. maculata is the object of targeted fisheries

Fig. 4. Grain size composition of sediment inside and outside shrimp burrows in dense and sparse seagrass beds. Error bars

represent standard error. Different letters above error bars indicate significant differences.

694 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 688-701, e46811, enero-diciembre 2022 (Publicado Set. 30, 2022)

(Ahyong et al., 2017; Abduho & Madjos, 2018;

Baigtu & Echem, 2018) and caught as by-catch

in other fisheries (Babu et al., 2022; Courtney

et al., 1999; Courtney et al., 2007). The zebra

mantis shrimps including L. maculata (and

perhaps L. sulcata as they are very similar) are

commonly found in big seafood restaurants in

Bali and Batam, Indonesia (Pers. Obs.). How-

ever, in general little is known of the social

behavior of lysiosquilloid mantis shrimps.

This study found that dense seagrass beds

harbor a higher abundance of zebra mantis

shrimp burrows than sparse seagrass beds. This

reinforces the view that seagrass beds are an

important habitat for economically important

animals (Nadiarti et al., 2015; Torre-Castro

et al., 2014), including mantis shrimps (Jay-

abarathi et al., 2013), with seagrass density

as an important factor with regards to the

abundance of these animals. Other studies have

reported higher faunal abundance in denser

seagrass beds, in general or in specific taxa

(e.g. Nadiarti et al., 2015; Nadiarti et al., 2021;

Pogoreutz et al., 2012; Vonk et al., 2008),

although the correlation can be weak for some

taxa (Priosambodo, 2015). Furthermore, sea-

grass community composition can also be an

important factor influencing faunal biodiver-

sity and abundance. For example, Nienhuis et

al. (1989) report that the density of selected

macrofauna was positively correlated with T.

hemprichii density but not with overall seagrass

density or the density of other seagrass species.

The seagrass species present at the study

site were the same as reported from seagrass

beds in this area by Nadiarti et al. (2021) who

considered the sea urchin and macroalgae

community composition were most likely influ-

enced by the identity and relative abundance

of the seagrass species present, in particu-

lar differences in canopy structure and hence

habitat niches available, as well as feeding

preferences and predator-prey relationships.

As mantis shrimps are carnivorous, predomi-

nantly piscivorous, ambush predators (deVries

et al., 2012), burrow surroundings are likely

an important factor with respect to the cap-

ture of prey. Habitat characteristics related to

seagrass community composition which might

influence mantis shrimp burrow site choice,

as well as individual growth and survival,

include the canopy structure (Pogoreutz et al.,

2012), root and rhizome structure (Kiswara

et al., 2009), and abundance of suitable prey

(Jackson et al., 2001).

Larger (diameter and/or depth) mantis

shrimp burrows were more common in the

denser seagrass beds and smaller burrows were

more common in sparse seagrass beds. This

pattern may indicate that larger mantis shrimps

prefer denser seagrass beds. A similar pattern

of larger individuals mostly found in denser

seagrass beds has been observed in seagrass

associated fishes (Nadiarti et al., 2015; Pogore-

utz et al., 2012). The higher abundance of man-

tis shrimp burrows in the dense seagrass beds

may be related to the structure of the seagrass

canopy and/or to root and rhizome structure.

As predators, mantis shrimps may prefer dense

seagrass cover which could provide a more

effective hiding place and make it easier to

ambush their prey than in a more exposed area

(sparse seagrass beds), as has been suggested

for predatory fishes (Schultz et al., 2009). The

denser and more structured seagrass beds could

provide better cover for the mantis shrimp to

ambush their prey from their burrow open-

ings; however, information about their activity

in their burrows and the surrounding areas is

extremely limited, calling for further studies on

mantis shrimp behavior and ecology.

Although the percentage cover was much

higher in the dense seagrass area, the seagrass

canopy was much taller in the sparse seagrass

area which was dominated by Enhalus acoroi-

des, the largest seagrass species found in Indo-

nesia. The thick rhizomes and long black bristly

cord-like roots of E. acoroides are closely inter-

woven and more deeply embedded into the

substrate than the root and rhizomes of other

seagrass species, and could be a challenge for

the zebra mantis shrimps when making their

burrows. In the dense seagrass area, the domi-

nant seagrasses were Cymodocea rotundata

and Thalassia hemprichii, both of which have

shorter canopies with smaller rhizomes and

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roots, among which it would most likely be

easier for zebra mantis shrimps to create large

(wide and/or deep) and stable burrows. On the

other hand, seagrass root systems can stabilize

sediment (Mcleod et al., 2011) and it is pos-

sible that, in stabilizing sediment, the roots may

also help to stabilize burrow walls. A more sta-

ble burrow will be better for the mantis shrimp,

since the burrow functions as a shelter, a place

for processing prey, and a safe home for mating

and for the guarding of eggs and larvae (Vetter

& Caldwell, 2015). The higher abundance of

mantis shrimp burrows in dense seagrass beds

compared to in sparse seagrass beds may be

influenced by these factors, although further

study is needed to test whether burrows are

more stable in dense seagrass areas, and to elu-

cidate the causal mechanisms for the observed

difference in burrow size and abundance.

The differences in burrow size are most

likely related to the size of the occupant,

indicating that burrows in the dense seagrass

tended to be occupied by larger mantis shrimps

compared to those in sparse seagrass. This may

indicate an ontogenetic shift in habitat prefer-

ence between the two seagrass habitats, but

could also be related to the species present.

Once possible explanation could be that juve-

niles are more abundant further offshore, and

will tend to move to the more favorable (pre-

sumably more productive) dense seagrass habi-

tat as they grow larger and are more capable

of constructing and defending their burrows.

The burrows not only serve as refuges and

for ambushing prey but also play a key role in

mantis shrimp reproduction (Mead & Caldwell,

2010; Matsuura & Hamano, 1984; Wortham-

Neal, 2002), as female mantis shrimps remain

in the burrow while brooding their eggs. The

juveniles of gonodactyloid smasher mantis

shrimps in the Caribbean live in deeper off-

shore reefs and migrate to more productive

inshore waters as they grow larger and are able

to fight for occupancy of the limited pre-exist-

ing cavities where reproduction can take place

(Reaka, 1987). However, in general, spearer

mantis shrimps are less aggressive than the

smasher mantis shrimps (Reaka and Manning,

1981). As they can construct their own bur-

rows, competition for burrows could be less

of an issue, although territorial behavior may

occur and could limit density and/or give rise to

competition over prime habitat. Further studies

are needed to elucidate the social behavior of

spearer mantis shrimps, including with respect

to their occupation of different habitats at

each life stage.

Although larger burrows (wider diam-

eter and deeper in depth) tend to occur in the

dense seagrass beds and small burrows in

sparse seagrass beds, there was no correlation

between diameter and depth. This indicates

that the inhabitants (mantis shrimp) of each

burrow construct dwellings of very different

shapes and sizes. Several studies show that

burrow dimensions and diameter size tend

to be correlated with the size of the inhabit-

ant (Atkinson et al., 1997; Mead & Caldwell,

2010; Matsuura & Hamano, 1984) as in some

gobies (Dinh et al., 2014), while size can also

be correlated with different species, as in fid-

dler crabs (Qureshi & Saher, 2012). Caldwell

& Dingle (1975) found different species of

mantis shrimp can create burrows of different

shapes and sizes, with burrow depth generally

proportional to the length of the mantis shrimp.

Therefore, the different burrow shapes may be

related to mantis shrimp size and/or species.

During our study we only found two zebra

mantis shrimp species (Lysiosquillina maculata

and L. sulcata); furthermore, mantis shrimps

caught by local fishers during the study period

were all identified as one of these two species.

However, were unable to verify which burrows

were inhabited by which species, and further

studies are needed to ascertain whether the

smaller burrows belong to smaller (possibly

juvenile or slow growing) zebra mantis shrimps

or belong to other mantis shrimp species.

The differences in burrow abundance and

size do not appear to be due to differences in

sediment between the dense and sparse sea-

grass beds. Despite significant differences in

some grain size classes both within and outside

the burrows, the overall substrate composi-

tion pattern within all burrows was similar in

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both seagrass bed types, with coarse and fine

particles more dominant than the medium

particles both in absolute terms and relative to

the composition of the surrounding substrate.

These data indicate that the mantis shrimps

were selective in the materials they used to

construct their burrows, in particular the inner

wall lining. Grain size selectivity in burrow

construction has been reported in several bur-

rowing marine invertebrates including crusta-

ceans (Sumida et al., 2020; Zorn et al., 2010).

The sediment outside and inside zebra

mantis shrimp burrows, in both sparse and

dense seagrass beds, was dominated by sand,

rubble and shell fragments (> 90 %) with the

remainder composed of silt and clay. This

means that the substrate of the zebra mantis

shrimp habitat in the study area is predomi-

nantly sandy, similar to the substrate character-

istics described by Nurdin et al., (2019) in the

same study area. Outside the burrows, coarse

sand (0.25-0.5 mm) was more abundant in

sparse than in dense seagrass beds; conversely,

medium sand (0.13-0.25 mm) was more abun-

dant in sparse than in dense seagrass beds.

These differences in sediment composition

were most probably due to the higher hydrody-

namic energy in sparse seagrass beds compared

with dense seagrass beds. Seagrasses reduce

hydrodynamic energy compared to unvegetated

areas (Contti Neto et al., 2022; Lanuru et al.,

2018; Potouroglou et al., 2017).

Furthermore, seagrasses have a differ-

ential effect on the transportation process for

particles of different sizes (Conley & Austin,

2017); in particular, the seagrass foliage may

reduce water movement and wave energy and

hence lower the effective transport rates of

coarser sediment particles with a higher set-

tling velocity, while turbulence generated by

the seagrass canopy can cause the resuspension

of finer particles. The higher proportion of

coarse particles in the area with sparse seagrass

compared to that with denser seagrass veg-

etation could seem counterintuitive. However,

this result is most likely related to the canopy

structure. The taller E. acoroides stands in the

sparse seagrass area most likely have a greater

effect in precipitating the deposition of coarse

particles; however, the higher velocities fur-

ther from the seabed and the greater distances

between shoots and patches may also enhance

turbulence which could facilitate the resuspen-

sion of finer particles and account for the lower

proportion of fine sediment. On the other hand,

the lower growing species in the dense seagrass

beds would be less likely to cause the pre-

cipitation of larger particles higher in the water

column; however, with very little exposed sub-

strate they are likely more effective in reducing

near-ground turbulence and retaining medium

and relatively fine particles. The proportion of

the largest grain sizes (> 1 mm and > 2 mm),

mostly comprised of coral rubble and shell

fragments, was similar in both seagrass bed

types, reflecting the proximity of these seagrass

beds to the coral reefs fringing the island and

that fact that they both harbor various shellfish.

The similarity in the grain size composi-

tion of particles lining the burrow walls of

zebra mantis shrimps in the dense or sparse

seagrass beds coupled with the marked dif-

ference between the burrow walls and the

composition of the surrounding substrate in

each seagrass area, indicates that the sediment

particles present around the burrow have a

limited influence on burrow wall construction,

with purposive selection (retention or rejection)

of excavated materials for use in burrow wall

construction. We did not observe how the zebra

mantis shrimps create their burrows, but we

assumed that burrow creation by zebra mantis

shrimps is similar to the way in which Squilla

empusa create their burrows, using their pleo-

pods for excavating and their maxillipeds for

basketing sediment (Mead & Minshall, 2012).

We based this assumption on morphological

similarities between the species, both of which

are spearers, sharing a similar raptorial append-

age structure; the dactyl of both species is

lined with sharp spines and the propodus is

also spined (Dingle & Caldwell, 1978). The

texture of the sediment from within the burrow

(burrow linings) was more solid than outside

the burrows, due to the mucus used in lining

the burrow wall. This mucus-reinforced lining

is essential in sandy sediment to support the

697

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 70: 688-701, e46811, enero-diciembre 2022 (Publicado Set. 30, 2022)

otherwise readily collapsing walls of the bur-

row (Papaspyrou et al., 2005).

Differences in burrow abundance and

size were not correlated with the water qual-

ity parameters measured (temperature, pH and

DO), as none of these parameters differed

significantly between sites. Although pH and

DO (but not temperature) were somewhat lower

within the burrows of mantis shrimp than out-

side the burrows (in both sparse and dense sea-

grass sites), the lack of statistical significance

was most likely related to the high between-

burrow variability reflected in the high stan-

dard error values (Table 1).

Although there was no significant correla-

tion between mantis shrimp burrow size and

seagrass density, the burrows were significantly

more abundant and larger in the dense seagrass

beds than the sparse seagrass beds. Seagrass

conservation is increasingly recognized as cru-

cial for climate mitigation, biodiversity protec-

tion, and food security (Cullen-Unsworth &

Unsworth, 2018). In particular, seagrass beds

provide vital habitat for various fauna with high

economic value, including mantis shrimps. The

results of this study highlight the importance

of conserving the quality (e. g. density) of sea-

grass ecosystems as well as their extent in order

to support seagrass-associated resources.

Ethical statement: the authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are

fully and clearly stated in the acknowledge-

ments section. A signed document has been

filed in the journal archives.

ACKNOWLEDGMENTS

The authors would like to express their

deep gratitude to the late Susan William for

input regarding the research design and to the

late Yusti for all her assistance. We thank Fitri

and Nita for assistance in the laboratory, and all

the students who helped in the field. We greatly

appreciate Dominic Kneer, Naomi Gardiner

and Laurence McCook for providing comments

on the original manuscript, including improv-

ing the English language usage. We thank the

anonymous reviewers for their constructive

comments and suggestions. We acknowledge

the contribution of Abigail Mary Moore, espe-

cially with regards to data analysis and manu-

script revision, and the contribution of C. B. de

los Santos for assistance with translating the

title and abstract into Spanish. This study was

funded under the Ministry of Research, Tech-

nology and Higher Education of the Republic

of Indonesia National Competitive Research

Grant Program (PTUPT grant No. 005/SP2H/

LT/DPRM/IV/2017 dated 20 April 2017).

RESUMEN

La densidad de los pastos marinos se correlaciona

con la abundancia y el tamaño de las madrigueras

de los camarones mantis cebra

(Stomatopoda: Lysiosquillidae)

Introducción: Los camarones mantis son organismos

ecológica y económicamente importantes en los ecosiste-

mas marinos. Sin embargo, aún falta información sobre su

hábitat, en particular sobre sus madrigueras.

Objetivo: Analizar cómo difieren las madrigueras de los

camarones mantis en su abundancia, tamaño, tamaño de

grano de los sedimentos y calidad del agua.

Métodos: Contamos las madrigueras en parcelas de 10 x

10 m2 al azar (diez parcelas por densidad) en pastos mari-

nos densos y poco densos, alrededor de la isla de Barrang

Lompo, Sulawesi del Sur, Indonesia.

Resultados: Se observaron dos especies de camarones

mantis: Lysiosquillina maculata y L. sulcata. El tamaño de

grano de las paredes de las madrigueras y los parámetros

de agua, tanto dentro y fuera de la madriguera no variaron

(P > 0.05). Tampoco hubo correlación entre la profundidad

y el diámetro de las madrigueras, tanto en praderas densas

(P > 0.05; r= 0.27), como no densas (P > 0.05; r= 0.33). Sin

embargo, las madrigueras más grandes tienden a aparecer

en las praderas densas, además había más madrigueras en

pastos densos (t-test, P < 0.05).

Conclusiones: Parece haber una preferencia por las prade-

ras marinas densas, especialmente en los camarones mantis

de mayor tamaño. La correlación entre la abundancia de

madrigueras de camarones y la densidad de pastos marinos

pone de manifiesto la importancia de conservar la calidad

del hábitat de los pastos, así como su extensión.

Palabras clave: Camarón mantis cebra; madrigueras;

pastos marinos; intermareal; isla de Barrang Lompo;

Indonesia.

698 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 70: 688-701, e46811, enero-diciembre 2022 (Publicado Set. 30, 2022)

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