Laguna (2023) 30: 61-70

ISSN 2185-2995

First record of the sea cucumber Holothuria (Selenkothuria)
erinaceus Semper, 1868 (Holothuroidea: Holothuriida:
Holothuriidae) from a Japanese mangrove forest
Shun Kawaida1, *, Yusuke Yamana2, Iwao Tanita3 , Kusuto Nanjo4

Abstract: The tropical epibenthic holothurian Holothuria (Selenkothuria) erinaceus Semper, 1868is reported for the first time; a single specimen has been collected from a subtropical mangrove
forest on Ishigaki Island, Yaeyama Islands, southern Japan. The external/internal morphologicalfeatures, including a calcareous ring and ossicles, are described and compared with literaturedescriptions of congeners. Holothuria (Selenkothuria) erinaceus Semper, 1868 has previously beenrecognized as a tropical species, distributed widely in the tropical Indian and West Pacific Oceans,including the Philippines and Fiji. Not only is the Ishigaki Island record the northernmost for the

species, but also the collection site is the first recorded instance of an epibenthic sea cucumber
inhabiting in a mangrove forest habitat.

Key words: Holothurians, Mangrove estuary, Morphology, Ossicles, Subtropical region(Lee, 2008). Kintz et al. (2013), for example, reported 156

Introduction

species of macrobenthic invertebrates from mangrove

Mangroves are salt-tolerant woody plants, widely

forests in Malaga Bay (136 km 2 ), Columbia, including

distributed in brackish intertidal zones in tropical andcrabs, shrimps, bivalves, gastropods and polychaetes.

subtropical regions (Kon et al., 2020). Mangrove forests

However, there appear to have been no publishedrecords of epifaunal sea cucumbers (Echinodermata:and adjacent tidal flats, defined as mangrove estuaries,
provide habitats for various macrobenthic invertebrates

Holothuroidea) occurring in a mangrove forest habitat,

1

Estuary Research Center, Shimane University, 1060 Nishikawatsu, Matsue, Shimane 690-8504, Japan.

2

Wakayama Prefectural Museum of Natural History, 370-1 Funo, Kainan, Wakayama 642-0001, Japan.

3

Yaeyama Field Station, Fisheries Technology Institute, Japan Fisheries Research and Education Agency, Ishigaki, Okinawa 907-0451, Japan.

4

Department of Applied Aquabiology, National Fisheries University, Japan Fisheries Research and Education Agency, Nagata–Honmachi,
Shimonoseki, Yamaguchi 759-6595, Japan.

* Corresponding Author

受付日：2023 年 6 月 15 日，受理日：2023 年 9 月 25 日，WEB 掲載日：2023 年 10 月 28 日

61

Shun Kawaida, Yusuke Yamana, Iwao Tanita , Kusuto Nanjo

Fig. 1 Map of the Fukido River mangrove estuary, Ishigaki Island, Ryukyu Islands, southern Japan.

The specimen of Holothuria (Selenkothuria) erinaceus was collected in the downstream section of
the river (indicated by white triangle).

despite the former being a common benthic faunal group,

Materials and Methods

and especially diverse in tropical and subtropical coastal
environments, including coral reefs, adjacent sandy

Locality and sampling method

Tanita and Yamada, 2019; Floren et al., 2021).

by hand from the muddy sediment surface shaded byin the subtropical region of southern Japan, due mainly

stylosa, a component of the subtropical mangrove forestal., 2004; Naruse, 2016). The present report describes(flowing into the East China Sea), located on northernbottoms and seagrass meadows (e.g., Pawson, 2007;

One specimen of the sea cucumber was collected

Tropical macrobenthic invertebrates are often found

an aerial prop root of the red mangrove Rhizophora

to larval transport by the Kuroshio Current (Komai et(24º29’12.3” N, 124º13’49.9” E) along the Fukido Rivera specimen of epibenthic sea cucumber of Holothuria

Ishigaki Island, Yaeyama Islands, southern Japan,(Selenkothuria) erinaceus, a tropical species occurring

between daytime (13:00–17:00) on 29 June 2022 (Fig. 1).

in the Indian and West Pacific Oceans, including the

The dense and undisturbed mangrove forest, extending

Philippines and Fiji (Semper, 1868), collected in aalong both sides of the river, was dominated by the

mangrove forest in southern Japan. Although several

black mangrove Bruguiera gymnorrhiza, with patches

other tropical and subtropical localities have beenof R. stylosa, with the water depth of the flat being

reported for the species, with the Philippines being theapproximately 1.5 m below mean high water spring tideprevious northernmost record (Semper, 1868; Rowe(MHWS) [prop roots inundated at high tide, and exposed

and Gates, 1995; Thandar and Samyn, 2004; Honey-

together with tidal flats at low tide (Fig. 2a, b)]. Sampling

Escandón et al., 2012), none have been reported fromof the sea cucumber was carried out with the permission

Japanese waters to date. Following a careful survey of thefrom the Fishery Division of the Okinawa Prefecture

morphological characteristics of the specimen, we report

Government.

the first record for Holothuria (Selenkothuria) erinaceus

Morphological observations

from Ishigaki Island in Japan, the present study

The specimen was fixed in 80% ethanol following

contributing to a better understanding of macrobenthic

anaesthetization, using magnesium sulfate (MgSO 4)

invertebrate species diversity in mangrove estuaries.

dissolved in sea water [concentration ca. 3.0% (w/w)].
62

First record of a tropical species of holothurid sea cucumber in Japan

Fig. 2 Habitat of Holothuria (Selenkothuria) erinaceus. Prop roots of the red mangrove Rhizophorastylosa (a). Tide level based on mean low water spring tide (MLWS) at Ishigaki Island on themeasurement date was indicated by white arrow. Holothuria (Selenkothuria) erinaceus on exposed
muddy substrate between prop roots (indicated by yellow arrow) (b).

The specimen was immersed in the latter for about three‘rods’ (i.e., rod-like ossicles, usually with distal crenation

were recorded by digital photographs, prior to fixation.

(multi-perforate flat ossicles). The specimen, glass-hours, after which live color and external characteristics

or teeth and few perforations); and ‘plates/endplates’

The specimen was dissected, exposing the tentacles

slides, and SEM stubs are deposited in the Invertebrate

and internal organs, including a calcareous ring, andCollection (INV) of the Wakayama Prefectural Museum

the eviscerated organs examined under a stereoscopicof Natural History (WMHN), Kainan, Wakayama, Japan.

dissecting microscope (SMZ1270, TREG-DSL32, Nikon
Japan Inc., Japan). A small piece of body-wall tissue

Results(ca. 2×2 mm) was sampled from both the mid-dorsal

Genus Holothuria Linnaeus, 1767and mid-ventral body wall, and several appendages

Subgenus Selenkothuria Deichmann, 1958

(five pedicels and five dorsal papillae, plus one dorso-

Diagnosis of subgenus (modified after Deichmann,interradial tentacle) extracted for observation of ossicle

morphology. Each tissue sample was dissolved using a1958) : Soft-skinned forms with numerous cylindricalCo., Ltd., Japan), and most of the extracted ossicles

sometimes arranged in three broad bands; dorsallycommercial bleach (Kitchen Bleach, Mitsuei Chemicalfeet in longitudinal bands forming distinct ventral sole,numerous minute papillae, not conspicuous, rarely

rinsed with distilled water, dried on glass slides, andmounted in polyester resin (RIGOLAC, Showadenko

forming low warts. Tentacles located at end of body, often

KK., Japan). The slides were examined and ossicles

bushy. Internal anatomy not remarkable (similar to manymeasured (to nearest µm) under a compound microscopeother subgenera); respiratory tree branching to both sidesof body cavity; some species with numerous stone canals(Optiphot XF-NT, Nikon Japan Inc., Japan). Several

in one or two tufts on both sides of dorsal mesenterium,

ossicles from each sample were rinsed with deionizedwater, dehydrated in 99% ethanol, and mounted onothers with single large stone canal attached to right side;

aluminum stubs using conductive tape (NEM Tape,usually one Polian vesicle; gonads apparent as divided

Nisshin EM Co., Ltd.). Samples were then dried at room

threads in tuft behind low calcareous ring; Cuverianorgans present in some forms. Table ossicles of external

temperature and observed using a scanning electron
microscope (SEM) (JCM-7000, JEOL Ltd., Japan).

body wall usually completely lacking, though vestiges

The examined ossicles were classified into two types:of tables sometimes found in young individuals of some63

Shun Kawaida, Yusuke Yamana, Iwao Tanita , Kusuto Nanjo

Fig. 3 Live specimen of Holothuria (Selenkothuria) erinaceus Semper, 1868 (WMNH-

INV-2022-366) collected from the Fukido River mangrove estuary. Dorsal side (a),ventral part (b), around oral disk and tentacles (c).

Clark and Rowe, 1971: 178. (distribution), pi. 28, fig. 5.

species. Ossicles of inner body wall consisting of small

Holothuria (Sel.) erinacea Thandar, 2007: 29–31, fig. 13.

rods or plates, either smooth or spinous. Ventral feetwith single large endplate and walls supported by rods

Original description (Semper, 1868: 91) : Twenty

or plates of same size as those in skin, rarely special
supporting rods or perforated plates present. Dorsal

short tentacles. Body cylindrical, slightly tapered towards

with more curved, shorter rods. Some species with rods

dense fine and long papillae on dorsal side, length ofboth ends. Numerous small feet on abdominal side. Very

appendages with or without reduced endplate, often

body 18–20 cm, maximum thickness 3.75 cm. Color ofpresent in cloacal retractor muscles and respiratory trees.

specimen plain grey-brown, anal field black. Specimen

Color dull gray, dark brown, olive-green or black, withcollected from Bohol Island, on shore, Viti Islands. Oneor without two rows of dark spots; anterior and posterior

Polian vesicle; 2 cm long stone canal, somewhat similarends darker.

to that of Holotliuria scabra Jäger. Whole reproductive

Holothuria (Selenkothuria) erinaceus Semper, 1868

organs of specimen 10 cm long in liquid, 2 cm from

[New Japanese name: Yaeyama-hirugi-namako] (Fig. 3)

calcareous ring, gonadal organs attached with very long

Synonymy

2–4 tubules; body of male specimen 5–6 cm shorter andsignificantly thinner than that of female described above.

Holothuria erinaceus Semper, 1868: 91, pi. 30, fig. 24.

Cuvierian tubules exceptionally small, in tuft at bottom

H. erinaceus var. pygmaea Semper, 1868: 91, pi. 30, fig.

of respiratory trees. Calcareous bodies of the specimens

23.

from Bohol Island lost completely due to acetic acid.

H. marenzelleri Ludwig, 1883: 167.

Specimen from Viti Islands no stool like ossicles,Η. (Halodeima) lubrica var. marenzelleri Panning, 1934:

numerous knotty rod like ossicles, extremely similar to47, fig. 41.

those of Holothuria glaberrima Selenka, but not unlikely

Η. (H.) lubrica var. glaberrima Panning, 1934: 47, fig. 42

both forms of ossicles observed together for specimens(part).

from other regions. C-shaped rod-like ossicles and plates

Selenk othuria erin a ceus Deich ma n n, 1958: 314

observed in pedicels, and rudimentary plates observed(synonymy), 315 (keys).

in body wall skin. Calcareous ring ribbon-like. Axillary

Holothuria (Selenkothuria) erinaceus Rowe, 1969: 135;64

First record of a tropical species of holothurid sea cucumber in Japan

F ig. 4 P r eser ve d sp e ci men of H ol o t h u r i a

Fig. 5 Elements of calcareous rings of Holothuria

INV-2022-366).

viewed from body cavity (anterior uppermost). Md,(Selenkothuria) erinaceus Semper, 1868 (WMNH-(Selenkothuria) erinaceus (WMNH-INV-2022-366)
mid-ventral radial zone.

cord in Cuvierian tubules not cellular, and scatteredprotected by narrow rhombus-shaped skin. Color of

Deutsch in Semper, 1868: 91 p.+ Plate XXX, f 24, mainly

ventral skin black-brown with three bands of scatteredliving specimen: tentacles semi-transparent yellow,

cells in fibrous connective tissue [originally described bybroad pale yellowish pedicels (greatest density < 10translated from original sentences to modern English bypedicels /cm 2), dorsal skin black-purple with dorsal

a language services provider (Crimson Interactive Japanyellowish papillae in about seven indistinct latitudinal

Co., Ltd.) with native German-speaking professionals].

lines (Fig. 3). Overall external morphology similar to thatof Holothuiria (Semperothuria) roseomaculata (reported

Type locality: Bohol Island, Philippines; Viti island,by Kerr, 2013). Color well maintained in preserved

Republic of Fiji (Semper, 1868).

specimen after six months (Fig. 4).

Species diagnosis (emended by Tortonese, 1980:

Internal morphology: Calcareous ring thick, with

pp. 105–106): Body becoming narrower anteriorly.

Uniformly brown, but clear ring around small buccal

solid radial and interradial plates, former convex, each

abdominal surface, much smaller and scattered oneach with sharp anterior prolongation (Fig. 5). Internally,

with one anterior and one posterior depression, latter

tentacles in 7 mm wide group. Podia crowded oneight tentacle ampullae, 7–10 mm in length; single

back, where skin much folded. Ossicles include rods

Polian vesicle, mid-ventral part close to calcareous ring,

only (generally more elongated than figured in original

cylindrical and expanding posteriorly (36 mm in length);description). Median axis apparent, somewhat recurved.

single stone canal, mid-dorsal part close to calcareous

Shape very variable, surface smooth, edge more or lessring attached to right side of mesentery, twisted and long

spiny, holes restricted at extremities or arranged in series(17 mm), with egg-shaped madreporite (1.2 mm in length;along one or both sides of axis. Larger rods forming

1.8 mm in breadth); Cuvierian tubules present, small,

perforated plates, smaller rods button-like; no true

tuft-like (eviscerated and could not be collected because

buttons or tables.

of the tubules sticking to the mangrove roots).

Material examined: One specimen, 176 mm long, 51

Ossicle morphology: Tentacle ossicles double-bladed

mm wide; body and eviscerated internal organs fixed andsaw-shaped rods in external (distal) part, short rods inpreserved in 80% ethanol (Registered number: WMNH-

proximal (basal) part (Fig. 6a). External spinous rods

INV-2022-366).

ranging between 147–205 µm in length, 8–11 µm in

breadth (at middle); proximal rods spinous, shorter thanExternal morphology: Oral disk slightly inclined toexternal rods, ranging between 56–70 µm in length, 9–11

ventral side. Tentacles 20 short (ca. 7.6 mm in preservedµm in breadth (at middle) (Table 1). Endplate situatedstate), forming single circle surrounding oral opening,
65

Shun Kawaida, Yusuke Yamana, Iwao Tanita , Kusuto Nanjo

Fig. 6 Kawaida, S., Yamana, Y., Tanita, I., Nanjo, K.

Fig. 6 SEM images of body ossicles of Holothuria (Selenkothuria)

erinaceus (WMNH-INV-2022-366). External and proximal rods fromlateral tentacle (a), endplate from pedicel (b), rods and/or plates frompapilla of anterior dorsal body wall (c), ventral body wall (d) and dorsal
body wall (e).

66

Table 1 Counts and measurements of ossicles extracted from a specimen of Holothuria
First record of a tropical species of holothurid sea cucumber in Japan(Selenkathuria) erinaceus.

Table 1 Counts and measurements of ossicles extracted from a
specimen of Holothuria (Selenkathuria) erinaceus.
Ossicle type and measurements (µm) n

Mean ± sd

Range

Tentacle external rods
Length4177 ± 25

147–205

Widtha4

10 ± 2

8–11

Length362 ± 7

56–70

Width

3

10 ± 1

9–11

Major axis

5640 ± 59581–709No. holesb

51024 ± 125

840–1158

Tentacle proximal rods

Pedicel endplate

Papilla rods and/or plates
Length

21 60 ± 8

48–76

Width

21 12 ± 4

5–19

Length15 60 ± 9

46–78

Width

15 15 ± 7

8–31

Length16 57 ± 10

41–77

Width

16 11 ± 4

5–18

Body rods and/or plates (ventral)

Body rods and/or plates (dorsal)

a Width of central part of stem.

b Estimated by doubling counts of half plate area.

on tips of pedicels, undulant, with numerous uneven

Lanka), Bay of Bengal, East Indies, northern Australia,

diameter ranging between 587–709 µm, with ca. 840–

Rowe, 1971), Australia (Rowe and Gates, 1995), Reunion

the Philippines and South Pacific Islands (Clark andperforations increasing and narrowing marginally,

and Comoros Island, France, Madagascar (Honey-

1158 holes (Fig. 6b, Table 1). Rod and plate ossicles

Escandón et al., 2012), and Ryukyu Islands, Japanabsent from pedicel wall; papillae and ventral/dorsal(present study).

body wall ossicles comprising derivatives of rod ossicles,
including rods, plates and intermediate forms (rod-likeplates), primarily exhibiting the shapes namely “Venetian

Discussion

mask-shape rods (Fig. 6c–e)”, ranging between 41–78 µm
in length, 5–31 µm in breadth (at middle) (Table 1).

The specimen collected was identified as Holothuria(Sel.) erinaceus; the ossicles were consistent with the

General distribution: Type localities (Semper, 1868);

original description and other taxonomic studies of the

Gulf Albay, Luzon Island, Bohol Island, Philippines,

species (Semper, 1868; Clark and Rowe, 1971). It differed

Viti Islands, Fiji, Port Mackay, Australia, and Singapore.

from phylogenetically related species (Honey-Escandón et

Also common from East Africa to Polynesia (Tortonese,al., 2012): H. (Sel.) bacilla (Cherbonnier, 1988), H. (Sem.)cinerascens (Ahmed et al., 2016), H. (Sel.) mactanensis1980), in tropical Indo-West Pacific Ocean (Rowe andGates, 1995; Thandar and Samyn, 2004), including east,(Tan Tiu, 1981), and H. (Sel.) parva (Thandar, 1977) inwest and southwest Pacific (Domantay, 1957), Ceylon (Srithe form of the body-wall ossicles, each predominantly
67

Shun Kawaida, Yusuke Yamana, Iwao Tanita , Kusuto Nanjo

having knobbed rods and thick/stock rods, tables andno instances of the species inhabiting mangrove forests

margin, and straight or irregular non-perforated spinous

reinforces that mangrove forests contribute to highrods, non-perforated sinuous rods with irregular spinose

reported to date. Accordingly, the finding of our study

rods, respectively, whereas H. (Sel.) erinaceus has only

biodiversity in coastal ecosystems by providing available

perforated short flat rods. Furthermore, other external/

habitats for various macrobenthic invertebrates, includinginternal morphological features reported in the present

holothurid sea cucumbers. The physical structure of

study completely matched the original description

mangrove vegetation reduces water movement (i.e.,(Semper, 1868), confirming the identity of the species.

wave and tidal current) energy and provides excellent

The present study not only represents the first record

shelter from coastal erosion for various macrobenthic

for H. (Sel.) erinaceus from Japan, but also extends theinvertebrates (Macintosh and Ashton, 2002). Such water

geographical distribution of the species approximately

movement would affect the distribution of some species

700 km north from the former northernmost recordof sea cucumbers such as Apostichopus japonicus and(Luzon Island, Philippines) (Semper, 1868). The status ofHolothuria scabra, which are typically found in sheltered

H. (Sel.) erinaceus in Japanese subtropical waters, either

areas with less wave and current action (Barkai, 1991;

well-established or a recent arrival, cannot be determined

Mercier et al., 2000). Furthermore, several studies haveat present, although a common inference can be drawn

revealed that large amounts of sediment organic matter

regarding northward distribution of marine organisms(comprised mainly mangrove detritus) are an importantaround Japan, due to the impact of the Kuroshio Current.

food source for some deposit feeders (Kawaida et al.,The latter flows northward near the eastern coast of

2019; Mackenzie et al., 2020). In general, holothurians

Taiwan, turns eastward to flow near the north side of theare considered to obtain nutrients from organic detritus

Ryukyu Islands, and subsequently flows near the southand sediments from substrate surface (Purcell et al.,coast of the Ohsumi Islands, and off Kyushu, Shikoku

2016), intestine of the specimen of H. (Sel.) erinaceusand Honshu (Asakura, 2021), with the potential to carry

being filled with fine mud probably due to feeding on

eggs and larvae of marine species from tropical regions

sediment organic matter in the mangrove forest floor.

to southern Japan and Honshu (Sassa et al., 2006; Sassa

These findings suggest that functions of mangrove forests

and Takasuka, 2020). Recent studies have shown that(i.e., shelter from wave/current impacts and organic-

various inland/estuarine brachyuran species with arich environments) are likely beneficial to H. (Sel.)pelagic larval phase, not previously recorded in Japan,

erinaceus. Further investigations on the relationships

have been transported from tropical areas such as the

between the spatial distribution of H. (Sel.) erinaceus

Philippines and India to the Yaeyama Islands, mainly

and abiotic environmental properties (e.g., salinity, propby the Kuroshio Current (Komai et al., 2004; Naruse,root densities, relative substrate elevation and sediment2016). Furthermore, some studies have indicated likely

characteristics) in mangrove forests would contribute toincreased habitat ranges of tropical species northwards,

a better understanding of the important role of mangrove

due to recent sea surface temperature rises related toestuaries in providing habitats for diverse macrobenthic

global warming (e.g., Bianchi and Morri, 1994; Hiscock

invertebrates.

et al., 2004; Crickenberger, 2014). Accordingly, larvaeof H. (Sel.) erinaceus may be transported northward

Acknowledgements

around Japan from tropical regions, although detailed
information on their pelagic duration and genetic

This work was funded by a Grant-in-Aid for Scientificstructure is necessary to clarify their population

Research (B) from the Japan Society for the Promotion

The H. (Sel.) erinaceus specimen was found on the

support from Dr. Francisco Alonso Solís Marín ofdynamics.

of Science (JSPS, No. 19H02976). We had enormous

muddy substrate in a mangrove forest. Although James

Colección Nacional de Equinodermos Instituto de(1982) reported that H. (Sel.) erinaceus inhabits mud

Ciencias del Mar y Limnología UNAM, who freely

flats in the supra-littoral zone, being gregarious in someshared his knowledge of Holothuria (Sel.) erinaceus.

places on the Andaman coast, India, there have been

An English language review by Dr. Graham Hardy was68First record of a tropical species of holothurid sea cucumber in Japanmuch appreciated. We also sincerely thank Mr. Masami

Floren, A.S., Hayashizaki, K.I., Putchakarn, S.,

Irie, Wakayama Prefectural Museum of Natural History,

Tuntiprapas, P. and Prathep, A. (2021) A review

Japan, for providing the equipment for SEM analysis.

of factors influencing the seagrass-sea cucumber

association in tropical seagrass meadows. Frontiers in
Marrne Science, 8: 696134.

References

Hiscock, K., Southward, A., Tittley, I. and Hawkins,

Ahmed, Q., Ali, M.Q. and Conand, C. (2016) New

S. (2004) Effects of changing temperature on

additions to the holothurian fauna of Pakistan:benthic marine life in Britain and Ireland. Aquatic
Conservation: Marine and Freshwater Ecosystems, 14:Holothuria verrucosa, Holothuria cinerascens
and Ohshimella ehrenbergii. SPC Beche-de-mer

333–362.

Information Bulletin, 36: 20–23.

Honey-Escandón, M., Laguarda-Figueras, A. and Solís-

Asakura, A. (2021) Crustaceans in changing climate:

Marín, F.A. (2012) Molecular phylogeny of the

global warming and invasion of tropical land hermit

subgenus Holothuria (Selenkothuria) Deichmann,crabs (Crustacea: Decapoda: Anomura: Coenobitidae)

1958 (Holothuroidea: Aspidochirotida). Zoologicalinto temperate area in Japan. Zoology, 145: 125893.

Journal of the Linnean Society, 165: 109–120.

Barkai, A. (1991) The effect of water movement on

James, D.B. (1982) Ecology of intertidal Echinoderms

the distribution and interaction of three holothurianof the Indian Seas. Journal of the Marine Biological

species on the South African west coast. Journal of

Association of India, 24: 124–129.

Experimental Marine Biology and Ecology, 153: 241–

Kawaida, S., Nanjo, K., Ohtsuchi, N., Kohno, H. andBianchi, C.N. and Morri, C. (1994) Southern species

the food utilization of mangrove estuarine crabs.

254.

Sano, M. (2019) Cellulose digestion abilities determine

in the Ligurian Sea (Northern Mediterranean) : new

Estuarine, Coastal and Shelf Science, 222: 43–52.

records and a review. Bollettino dei musei e degli

Kerr, A.M. (2013) Holothuria (Semperothuria)

istituti biologici dell' Università di Genova, 58: 181–

roseomaculata n. sp. (Aspidochirotida: Holothuriidae),197.

a coral-reef inhabiting sea cucumber from the western

Cherbonnier, G. (1988) Echinodermes : Holothurides.

Pacific Ocean. Zootaxa, 3641: 384–394.

Faune de Madagascar, 70: 1–292.

Kintz, J.R.C., Londoño-Cruz, E., Mejía-Ladino, L.M.,

Clark, A.M. and Rowe, F.W.E. (1971) Monograph

Herrera-Orozco, L., Satizabal, C.A. and Uribe-of shallow-water Indo-West Pacific Echinoderms.

Castañeda, N. (2013) Environmental issues of a marine

Trustees of the British Museum (Natural History),

protected area in a tectonic estuary in the tropicalLondon, 238 pp.+31 Plates.

eastern Pacific : Uramba (Malaga Bay Colombia):

Crickenberger, S. (2014) Range limits, range shifts,

context, biodiversity, threats and challenges. Journal ofand lower thermal tolerance in the tropical barnacle

Water Research and Protection, 5: 1037–1047.

Megabalanus coccopoma. PhD thesis. Clemson

Komai, T., Nagai, T., Yogi, A., Naruse, T., Fujita, Y. andDeichmann, E. (1958) The Holothuroidea collected by(Crustacea: Decapoda: Brachyura) from Japan, with

University, Clemson, SC, USA.

Shokita, S. (2004) New records of four grapsoid crabs

the Velero III and IV during the years 1932 to 1954.

notes on four rare species. Natural History Research, 8:Part. II Aspidochirota. The University of Southern

33–63.

California Publications Allan Hancock PacificKon, K., Shimanaga, M. and Horinouchi, M. (2020)

Domantay, J.S. (1957) The zoogeographical distribution

Marine Life, (eds.) Inaba, K. and Hall-Spencer, J.M.

Expeditions, 11: 253–348.

Marine Ecology: Intertidal/Littoral Zone. In: Japaneseof the Indo-Pacific littoral Holothurioidea. In:

pp. 241–254. Springer, Singapore.

Proceedings of the Eighth Pacific Science Congress

Lee, S.Y. (2008) Mangrove macrobenthos: assemblages,of the Pacific Science Association. National Researchservices, and linkages. Journal of Sea Research, 59:

Council of the Philippines, Quezon, Philippines, pp.

16–29.

417–455.

Linnaeus, C. (1767) Systema naturae per regna tria
69

Shun Kawaida, Yusuke Yamana, Iwao Tanita , Kusuto Nanjo

naturae: secundum classes, ordines, genera, species,

Sassa, C. and Takasuka, A. (2020) Distribution and

Ed. 12. 1., Regnum Animale. 1 & 2. Laurentii Salvii,

Myctophidae) on the onshore side of the Kuroshio

cum characteribus, differentiis, synonymis, locis.

feeding of Myctophum orientale juveniles (Teleostei:

Holmiae (Stockholm), pp. 533–1327. (in Latin)

off Japan. Deep-Sea Research Part I: Oceanographic

Ludwig, H. (1883) Verzeichniss der Holothurien

Research Papers, 162: 103318.

des Kieler Museums. In: Festschrift zur Feier desSemper, C. (1867–1868) Holothurien. In: Reisen im

der Oberhessischen Gesellschaft für Natur- undpp., 40 pls. [1867 = pp. 1–70, pls. 1–15; 1867/(1868)

fünfzigjährigen Bestehens der Gesellschaft. Bericht

Archipel der Philippinen, (ed.) Semper, C. iv + 288

Heilkunde, 22: 155–176. (in German)= pp. 71–100, pls. 16−25; 1868 = pp. 101–288, pls.

Macintosh, D.J. and Ashton, E.C. (2002) A review of

26–40]. Zweiter Theil, Wissenschaftliche Resultate,mangrove biodiversity conservation and management.

Erster Band, Leipzig: W. Engelmann. (in German)

Centre for Tropical Ecosystems Research, University

Tanita, I. and Yamada, H. (2019) Distribution of sea

Mackenzie, R.A., Cormier, N. and Demopoulos, A.W.

coral reef lagoons and adjacent waters around Ishigaki

of Aarhus, Denmark, pp. 1–71.

cucumbers in relation to sediment characteristics in(2020) Estimating the value of mangrove leaf litter inIsland, southern Japan. Marine Ecology, 40: e12564.

sesarmid crab diets : the importance of fractionation

Tan Tiu, A.S. (1981) The intertidal holothurian fauna

Mercier, A., Battaglene, S.C. and Hamel, J.-F. (2000)

neighboring islands, central Philippines. Philippine

factors. Bulletin of Marine Science, 96: 501–519.

(Echinodermata: Holothuroidea) of Mactan and the

Periodic movement, recruitment and size-related

Scientist, 18: 45–119.

distribution of the sea cucumber Holothuria scabra inThandar, A.S. (1977) Descriptions of two new species

Solomon Islands. Hydrobiologia, 440: 81–100.

of Holothuroidea from the East Coast of South Africa.

Naruse, T. (2016) Characteristics and recent findings

Annals of the Natal Museum, 23: 57–66.

of the invertebrates of inland waters in the Ryukyu

Thandar, A.S. (2007) Additions to the aspidochirotid,

Islands. Japanese Journal of Limnology, 77: 203–209.

molpadid and apodid holothuroids (Echinodermata:(in Japanese)

Holothuroidea) from the east coast of southern Africa,

Panning, A. (1934) Die Gattung Holothuria. Mitteilungenwith descriptions of new species. Zootaxa, 1414: 1–62.

aus dem Zoologischen Staatsinstitut und Zoologischen

Thandar, A.S. and Samyn, Y. (2004) Shallow-water

Pawson, D.L. (2007) Phylum Echinodermata. Zootaxa,

biodiversity and biogeography of the subtropical east

Museum in Hamburg, 45: 24–50. (in German)

holothuroid (Echinodermata: Holothuroidea)

1668: 749–764.

coast of South Africa. In: Echinoderms München, andPurcell, S.W., Conand, C., Uthicke, S. and Byrne, M.

Proceedings of the 11th International Echinoderm(2016) Ecological roles of exploited sea cucumbers.

Conference, (eds.) Heizeller, T. and Nebelsick, J. pp.

Oceanography and Marine Biology: An Annual

253–260. A.A. Balkema, Rotterdam.

Review, 54: 367–386.

Tortonese, E. (1980) Researches on the coast of Somalia.

R o w e , F. W. E . ( 1 9 6 9 ) A r e v i e w o f t h e f a m i l y

Littoral Echinodermata. Monitore Zoologico Italiano.

Holothuriidae (Holothurioidea: Aspidochirotida).

Supplemento, 13: 99–139.

Bulletin of the British Museum (Natual Histroy), 18:
117–170.

Rowe, F.W.E. and Gates, J. (1995) Echinodermata. In:
Zoological Catalogue of Australia, (ed.) Wells, A. 510
pp. CSIRO Australia, Melbourne.

Sassa, C., Konishi, Y. and Mori, K. (2006) Distributionof jack mackerel (Trachurus japonicus) larvae andjuveniles in the East China Sea, with special reference
to the larval transport by the Kuroshio Current.
Fisheries Oceanography, 15: 508–518.

70