Hermit crabs (Decapoda: Anomura: Coenobitidae, Calcinidae, Diogenidae, Paguridae) from Halmahera Island, North Maluku, Indonesia

Abstract

A total of 340 specimens of hermit crabs belonging to 34 species, included in four families and 10 genera were collected from the northern coast of Halmahera, in North Maluku, Indonesia. Two species are reported for the first time from Indonesia: Coenobita pseudorugosus Nakasone, 1988, and Trichopagurus macrochela Komai and Osawa, 2005. The color in life for Diogenes spinicarpus Rahayu and Forest, 1995, Diogenes foresti Rahayu and Hortle, 2002, and Clibanarius bistriatus Rahayu and Forest, 1993 is also reported for the first time. A list of hermit crabs found in northern Halmahera, including its habitat and distribution is presented here.

Keywords:
Anomura; Halmahera; new record; shallow waters; taxonomy.

INTRODUCTION

Hermit crabs are the most ubiquitous organisms in shallow waters of tropical areas. In Indonesian waters, they are relatively well studied, with 205 species recorded from shallow intertidal to deep waters (Rahayu and Wahyudi, 2008; Rahayu, 2021; Rahayu and Pratiwi, 2022). The study of littoral hermit crabs in Indonesia was started in the 19^th century, with the specimens mainly collected from Maluku islands (cf. Rahayu and Wahyudi, 2008). However, the occurrence of hermit crabs from Halmahera Island in the northern part of Maluku, Indonesia, was only reported by Rahayu and Forest (1993; 1995).

Halmahera Island is one of the largest islands in Maluku, located in the northern part of the archipelago, the western side facing the Pacific Ocean, while the eastern part is adjacent to the Maluku Sea. A survey of the diverse habitats of littoral areas, including reef flat, coral reef, seagrass beds, gravels, boulders, and rocky shores, mangrove, sandy, and muddy estuarine in the northern part of Halmahera and the neighbouring islands has resulted in a collection of 34 species of hermit crabs belonging to four families and ten genera. All these species are known to have a wide distribution in the Indo-Pacific, except for Coenobita pseudorugosusNakasone, 1988, which was known only from Cebu, the Philippines, Diogenes forestiRahayu and Hortle, 2002, known only from Papua, Indonesia, and Trichopagurus macrochelaKomai and Osawa, 2005 known from Ryukyu, Japan, Palau, and the Philippines. Considering that Halmahera is the largest island in northern Maluku with various habitats, very little is known about its marine crustaceans. This study was intended to collect littoral crustacean data, the hermit crabs listed in this paper, with information on their habitat and distribution in Indonesia, are one of several reports on crustaceans from Halmahera to come.

MATERIALS AND METHODS

Hermit crabs were collected by hand during low tide from intertidal areas from diverse habitats on the northeast coast of Halmahera: Gamsungi, 01°43’45.9”N 128°00’41.3”E, 8 August 2022; Galela, 01°47’53.2”N 127°53’1.3”E, 8 August 2022; Pulau Tagalaya, 01°43’33.9”N 128°03’58.2”E, 9 August 2022; Pulau Kakara, 01°44’34.4”N 128°02’24.0 E, 10 August 2022; Pulau Kumo, 01°43’59.9”N 128°01’23,6”E, 11 August 2022; Pulau Tolonuo, 01°47’14.0”N 127°59’55.4”E, 12 August 2022 (Figs. 1, 2).

Figure 1.
(A) Map of Indonesia, showing Halmahera Island. (B) Halmahera Island. (C) Collection sites at North Halmahera. (1) Gamsungi. (2) Pulau Tagalaya. (3) Pulau Kakara. (4) Pulau Tolonuo. (5) Pulau Kumo. (6) Galela.

Figure 2.
(1, 2) Gamsungi: mud and mangrove habitat. (3, 4) Pulau Tagalaya, coral reef and mangrove habitat. (5, 6) Pulau Kumo, rocky shore and seagrass bed. (7) Pulau Tolonuo, rocky shore. (8) Galela, estuarine.

The specimens collected were photographed, preserved in 70% ethanol, and deposited in the Bogor Zoology Museum (MZB) of the National Research and Innovation Agency (BRIN) (formerly Indonesian Institute of Sciences-LIPI) in Cibinong, Indonesia. Animal size, in mm, indicated by shield length, was measured from the tip of the rostrum to the midpoint of the posterior margin of the shield. Ambulatory legs are the second and third pereopods. Color in life were given in detail for the specimens photographed in preservative.

Comparative material: Clibanarius bistriatusRahayu and Forest, 1993, 1 female, 2.1 mm, Teluk Kodek, Lombok Island, Indonesia, 4 June 2023; Diogenes spinicarpus Rahayu and Forest, 1995, 2 males, 1.3 mm, 1.8 mm, Kuta, Lombok Island, Indonesia, 18 June 2023.

RESULTS

A total of 340 specimens of hermit crabs belonging to 34 species of hermit crabs, included in four families and 10 genera, were collected (Tab. 1). Family Coenobitidae with one genus and two species, family Calcinidae with three genera and 10 species, family Diogenidae with three genera and 16 species, and family Paguridae with three genera and six species.

Table 1.
List of hermit crabs collected from Halmahera Island, Indonesia.

Family/Species	Material examined
Family Coenobitidae
Coenobita rugosus H. Milne Edwards, 1837	2 males, 3.4-3.9 mm, 6 females, 2.9-4.9 mm (MZB Cru 5501), Pulau Tolonuo; 1 male, 3.8 mm (MZB Cru 5502), Pulau Kakara; 2 males, 2.9-3.4 mm, 24 females, 3.4-4.8 mm (MZB Cru 5503), Pulau Kumo; 8 males, 3.1-5.3 mm, 18 females, 2.3-7.3 mm, 1 ovigerous female, 4.9 mm (MZB Cru 5504), Pulau Tagalaya.
Coenobita pseudorugosus Nakasone, 1988	3 males, 5.3-5.9 mm, 2 females, 3.9-4 mm, 3 ovigerous females 4.7-5 mm (MZB Cru 5505), Pulau Kakara; 1 male, 4.1 mm (MZB Cru 5506), Pulau Tagalaya.
Family Calcinidae
Calcinus gaimardii (H. Milne Edwards,1848)	2 males, 3.5-4.9 mm (MZB Cru 5507), Pulau Kumo.
Calcinus laevimanus (Randall, 1840)	6 males,1.6-4.7 mm, 4 females 2.0-3.1 mm, 1 ovigerous female (MZB Cru 5508), Pulau Tolonuo; 2 males, 2.2-4.1 mm (MZB Cru 5509), Pulau Kakara.
Calcinus latens (Randall, 1840)	1 male, 4.4 mm (MZB Cru 5510), Pulau Tolonuo; 2 females, 4-5.5 mm (MZB Cru 5511), Pulau Kakara; 1 male, 4.0 mm (MZB Cru 5512), Pulau Kumo; 2 males, 3.5-4.9 mm (MZB Cru 5513), Pulau Tagalaya.
Calcinus morgani Rahayu and Forest, 1999	1 male, 1.8 mm (MZB Cru 5514), Pulau Kakara.
Ciliopagurus strigatus (Herbst, 1804)	1 male, 4.7 mm (MZB Cru 5515), Pulau Tagalaya.
Dardanus deformis (H. Milne Edwards, 1836)	2 females, 9.1-14.4 mm (MZB 5516), Pulau Kakara; 1 female, 6.2 mm (MZB Cru 5517), Pulau Tagalaya.
Dardanus guttatus (Olivier, 1812)	1 male, 4.9 mm (MZB Cru 5518), Pulau Kakara.
Dardanus lagopodes (Forskål, 1775)	1 male, 5.5 mm (MZB Cru 5519), PulauTolonuo; 1 male, 7.1 mm (MZB Cru 5520), Pulau Kumo.
Dardanus megistos (Herbst, 1804)	1 male, 10.7 mm (MZB Cru 5521), Pulau Kakara; 1 female, 9.6 mm (MZB Cru 5522), Pulau Tagalaya.
Dardanus scutellatus (H. Milne Edwards, 1848)	1male, 3.6 mm (MZB Cru 5523), Pulau Tolonuo.
Family Diogenidae
Clibanarius ambonensis Rahayu and Forest, 1993	1 male, 5.3 mm, 2 females, 4.2 mm, 1 ovigerous female, 4.9 mm (MZB Cru 5524), Galela.
Clibanarius cruentatus (H. Milne Edwards, 1848)	3 females, 2.2-4.9 mm (MZB Cru 5525), Pulau Kumo.
Clibanarius cf. demaniBuitendijk, 1937	1 male, 2.7 mm (MZB Cru 5526), Pulau Kumo.
Clibanarius englaucus Ball and Haig, 1972	6 males, 1.5-2.0 mm, 1 ovigerous female, 1.3 mm (MZB Cru 5527), Pulau Tolonuo; 5 males, 1.8-2.4 mm (MZB Cru 5528), Pulau Kakara; 3 males, 1.8-2.4 mm (MZB Cru 5529), Pulau Kumo.
Clibanarius eurysternus (Hilgendorf, 1879)	4 males, 2.4-3.5 mm, 1 female, 2.5 mm (MZB Cru 5530), Pulau Tolonuo; 2 males, 1.5-3.5 mm (MZB Cru 5531) Pulau Kakara; 1 male, 2.7 mm (MZB Cru 5532), Pulau Tagalaya.
Clibanarius longitarsus (De Haan, 1849)	1 male, 2,7 mm, 2 ovigerous females, 2,4 mm (MZB Cru 5534), Gamsungi.
Clibanarius merguiensis De Man. 1888	1 male, 2.4 mm (MZB Cru 5535), Gamsungi; 8 males. 1.6-2.4 mm (MZB Cru 5536), Pulau Tolonuo; 9 males, 2.0-3.3 mm, 1 female, 2.5 mm, 3 ovigerous females, 2.0-2.2 mm (MZB Cru 5537), Pulau Kakara; 14 males, 1.8-4.2 mm, 1 female, 2.5 mm, 2 ovigerous females, 2.7-2.7 mm (MZB Cru 5538), Pulau Kumo.
Clibanarius snelliusi Buitendijk, 1937	1 male, 3.3 mm (MZB Cru 5539), Pulau Kumo.
Clibanarius striolatus Dana, 1852	10 males, 2.4-6.4 mm, 1 ovigerous female, 2 mm (MZB Cru 5540), Gamsungi; 11 males, 1.3-6.4 mm, 2 females, 1.8-4.0 mm, 2 ovigerous females, 1.4-3.3 mm (MZB Cru 5541), Pulau Tolonuo; 3 males, 4.0-5.5 mm, 1 female, 4.4 mm, 2 ovigerous females, 3.6-4.5 mm (MZB Cru 5542), Pulau Kakara; 1 male, 3.3 mm (MZB Cru 5543), Pulau Kumo.
Clibanarius virescens (Krauss, 1843)	2 males, 3.1-4.1 mm (MZB Cru 5544), Pulau Tolonuo; 3 males, 2.0-2.5 mm (MZB Cru 5545), Pulau Kakara.
Diogenes avarus Heller, 1865	1 male, 1.5 mm (MZB Cru 5546), Pulau Kakara. 1 male, 3.5 mm (MZB Cru 5533), Gamsungi.
Diogenes foresti Rahayu and Hortle, 2002	12 males, 1.3-3.5 mm, 1 ovigerous female, 1.5 mm (MZB Cru 5547), Gamsungi; 1 male, 2.8 mm, 1 ovigerous female, 2.5 mm (MZB Cru 5548), Pulau Tolonuo; 2 males, 1.8-2.9 mm (MZB Cru 5549), Pulau Kumo.
Diogenes pallescens Whitelegge, 1897	2 ovigerous females, 1.3-1.6 mm (MZB Cru 5550), Pulau Tolonuo; 4 males,1.1-1.6 mm, 5 ovigerous females, 1.0-1.8 mm (MZB Cru 5551), Pulau Kakara; 1 male, 1.3 mm (MZB Cru 5552), Pulau Kakara; 3 males, 1.5-1.6 mm, 1 ovigerous female, 1,6 mm (MZB Cru 5553), Pulau Tagalaya.
Diogenes spinicarpus Rahayu and Forest, 1995	7 males, 1.3-2.2 mm, 7 ovigerous females, 1.3-2.4 mm (MZB Cru 5554), Gamsungi; 2 males, 1.8-2.0 mm, 1 ovigerous female, 2.0 mm (MZB Cru 554355), Pulau Tolonuo.
Diogenes tumidus Rahayu and Forest, 1995	2 ovigerous females, 1.8 mm (MZB Cru 5556), Gamsungi; 5 males, 1.3-2.4 mm, 2 ovigerous females, 1.7-2.0 mm (MZB Cru 5557), Pulau Kakara.
Pseudopaguristes monoporus (Morgan, 1987)	1 male, 1.3 mm (MZB Cru 5558), Pulau Tolonuo; 7 males, 1.6-4.7 mm, 5 ovigerous females, 2.2-2.9 mm (MZB Cru 5559), Pulau Kakara; 5 males, 1.3-3.5 mm, 3 ovigerous females, 2.4-2.5 mm (MZB Cru 5560), Pulau Kumo; 1 male, 1.8 mm (MZB Cru 5561), Pulau Tagalaya.
Family Paguridae
Pagurus fungiformis Komai and Rahayu, 2004	5 males, 5 ovigerous females, 8 individuals in shell (MZB Cru 5562), Pulau Kakara; 7 ovigerous females1.6-2.4 mm (MZB Cru 5563), Pulau Kumo; 1 male, 2.9 mm (MZB Cru 5564), Pulau Kumo.
Pagurus hirtimanus (Miers, 1880)	2 males, 4.2-4.5 mm, 3 ovigerous females, 4.0-4.5 mm (MZB Cru 5565), Pulau Tolonuo; 1 male, 3.1 mm (MZB Cru 5566), Pulau Kakara; 5 males, 2.7-5.5 mm, 1 female, 4.0 mm, 3 ovigerous females, 4.0-7.5 mm (MZB Cru 5567), Pulau Kumo; 6 males, 2.7-4.7 mm (MZB Cru 5568), Pulau Tagalaya.
Pagurus moluccensis Haig and Ball, 1988	1 male, 1.8 mm (MZB Cru 5569), Pulau Kumo.
Pagurixus anceps (Forest, 1954)	1 male, 1.3 mm (MZB Cru 5570), Pulau Tolonuo.
Pagurixus haigae Komaiand Osawa, 2007	2 males, 1.4-2.0 mm (MZB Cru 5571), Pulau Kakara.
Trichopagurus macrochela Komai and Osawa, 2005	1 ovigerous female 1.8 mm (MZB Cru 5572), Pulau Tagalaya.

SYSTEMATICS

Family Coenobitidae Dana, 1851

Genus Coenobita Latreille, 1829

Coenobita rugosus H. Milne Edwards, 1837

(Figs. 3A‒C, 4A)

Habitat. Upper intertidal, sandy beach with gravel, dead coral, and leaf litter and vegetation.

Distribution. Wide distribution in the Indo-West Pacific, from the East coast of Africa to Tuamotu archipelago. Indonesia: Sulawesi and Maluku.

Remarks. Coenobita rugosus was recorded from most of the islands in Maluku but not to date from Halmahera (see Buitendijk, 1937; Haig and Ball, 1988). The presence of stridulatory apparatus on the left cheliped palm, and the shield and cheliped whitish brown with streaks of dark brown or reddish brown (Figs. 3A, 4A) easily separate this species from most of its congeners. Differences with C. pseudorugosus are discussed under the species below. Compared to the specimen examined by Nakasone (1988), which has the right male sexual tube slightly more slender than the left, the specimens of this study have the right and left male sexual tube approximately similar (Nakasone, 1988: fig. 2G; Fig. 3C in this study). This may be due to the fact that the specimens in this study are smaller than those examined by Nakasone (1988) and the sexual tube is not fully developed yet.

Figure 3.
(A-C) Coenobita rugosus H. Milne Edwards, 1837, male, 4.5 mm (MZB Cru 5504). (D-F) Coenobita pseudorugosusNakasone, 1988, male 5.9 mm (MZB Cru 5505). (A, D) left cheliped. (B, E) Left third pereopod. (C, F) Sternite and coxa of male fifth pereopod. Photo in preservative.

Coenobita pseudorugosus Nakasone, 1988

(Figs. 3D‒F, 4B)

Habitat. Upper intertidal, sandy beach with gravel, dead coral, and leaf litter and vegetation.

Distribution. Cebu, Philippines. Indonesia: North Halmahera, this is the first record in Indonesia.

Remarks. This is the first record of the species after its description by Nakasone in 1988. Among the species with stridulatory apparatus on the left cheliped, this species mostly resembles C. rugosus. The differences of these two species agree well with Nakasone’s (1988) observation: the lower margin of the palm of the left cheliped is straight in the distal half in C. pseudorugosus (Fig. 3D), versus more convex in C. rugosus (Fig. 3A); right male sexual tube longer and more slender than the left in C. pseudorugosus (Fig. 3F), versus left and right male sexual tube similar, or right very slightly longer (Fig. 3C). Furthermore, in C. rugosus the dark brown patch on the left cheliped palm is clearly indicated even after preservation (Fig. 3A), while in C. pseudorugosus it is blurred (Fig. 3D).

Family Calcinidae Fraajie, Van Bakel and Jagt, 2017

Genus Calcinus Dana 1851

Calcinus gaimardii (H. Milne Edwards, 1848 )

(Fig. 4C)

Habitat. Coral reef, intertidal to 3 m.

Distribution. Wide distribution in the Indo-Pacific. Indonesia: Maluku and Lombok.

Remarks. The specimens in this study agree well with the color description of the species by Morgan (1991) and Rahayu and Forest (1999), i.e., the entire or most of the surface of the shield is dark brown, the ocular peduncles are brown on the proximal three-quarters but the distal quarter is blue.

Figure 4.
(A) Coenobita rugosus H. Milne Edwards, 1937, female 7.3 mm (MZB Cru 5504). (B) Coenobita pseudorugosusNakasone, 1988, male, 4.1 mm (MZB Cru 5506). (C) Calcinus gaimardii (H. Milne Edwards, 1848), male, 4.9 mm (MZB Cru 5507). (D) Calcinus laevimanus (Randall, 1840), male 4.7 mm (MZB Cru 5508). (E) Calcinus latens (Randall, 1840), male, 3.5 mm (MZB Cru 5513). (F) Calcinus morganiRahayu and Forest, 1999 (photo in preservative), male, 1.8 mm (MZB Cru 5514). Live coloration unless indicated otherwise.

Calcinus laevimanus ( Randall, 1840 )

(Fig. 4D)

Habitat. Intertidal, on sandy muddy bottom with gravel and coral rubbles.

Distribution. Wide distribution in the Indo-West Pacific, from the East coast of Africa to Hawaii and Tuamotu archipelago. Indonesia: Sulawesi, Maluku, and Lombok.

Remarks. This is a quite common species in gravel and sandy beach in Indonesian waters. Usually found under rocks or boulders during low tide. In other places of the Indo-West Pacific, C. laevimanus is also found in hard substrate of upper littoral zone, rocky platform, shallow pools and towards upper part of shore with sparse algal growth (McLaughlin et al, 2007; Malay et al., 2018). The species is easily recognized by the black and white color of the left cheliped and the blue band on the ocular peduncles. This coloration persists in preservative.

Calcinus latens ( Randall, 1840 )

(Fig. 4E)

Habitat. Intertidal, on seagrass bed, dead coral, and sandy mud substrates.

Distribution. Wide distribution in the Indo-Pacific. Indonesia: Sulawesi, Maluku and Lombok.

Remarks. This is a very common species of the intertidal, often found aggregated under large boulders during low tide. Calcinus latens is easily recognized in life by the bluish-green color of the shield and chelipeds, the ambulatory legs dactylus with dark brown or purple stripes proximally, the propodus is light brown, and carpus and merus are bluish green.

Calcinus morgani Rahayu and Forest, 1999

(Fig. 4F)

Habitat. Coral Reef, intertidal to 3 m.

Distribution. Wide distribution in the Indo-Pacific. Indonesia: Maluku, Sulawesi, and Lombok.

Color in life. Shield grey white or cream; ocular peduncles dark brown proximally, distal half blue with narrow dark ring close to cornea. Antennular peduncles dark brown, flagella yellow; antennal peduncles and flagella yellow-orange. Cheliped dark brown; ambulatory legs brown to dark brown, dactyl, propodus, carpus, and merus sometimes with dark brown spots (after Komai, 2004).

Remarks. Rahayu and Forest (1999) described C. areolatus, a species that morphologically closely resembles C. morgani but with different coloration, i.e., the ambulatory legs dactylus and propodus white with small red dots on the entire surface. Komai (2004) documented that C. areolatus was based on juvenile of C. morgani therefore the morphological character and coloration were not fully attained. The specimen in this study is small in size and concurred with Komai’s observation. Calcinus gaimardii from Maluku reported by Buitendijk (1937) is possibly a juvenile of C. morgani considering that the color she gave resembles that of C. areolatus.

Genus Ciliopagurus Forest, 1995

Ciliopagurus strigatus (Herbst, 1804)

(Fig. 5A)

Habitat. Rubble beach, coral reef, intertidal to 10 m.

Distribution. Wide distribution in the Indo-Pacific. Indonesia: Maluku and Lombok.

Remarks. Among the 19 species included in the genus Ciliopagurus worldwide, only three species are known to occur in Indonesian waters so far: C. strigatus lives in intertidal area, C. krempfi (Forest, 1952), and C. macrolepis Forest, 1995 inhabit deeper waters (70‒85m) (Forest, 1995). Ciliopagurus strigatus is easily recognized by the red and yellow stripes that are embellished with short setae on the chelipeds and ambulatory legs.

Figure 5.
(A) Ciliopagurus strigatus (Herbst, 1804), male, 4.7 mm (MZB Cru 5515). (B) Dardanus guttatus (Olivier, 1812), male, 4.9 mm (MZB Cru 5518). (C) Dardanus deformis (H. Milne Edwards, 1848) female, 14.4 mm (MZB Cru 5516). (D) Dardanus deformis (H. Milne Edwards, 1848), female, 9.1 mm (MZB Cru 5516). (E, F) Dardanus megistos (Herbst, 1804), male, 10.7 mm (MZB Cru 5521). Live coloration unless indicated otherwise.

Genus Dardanus Paul’son 1875

Dardanus deformis (H. Milne Edwards, 1836 )

(Fig. 5C, D)

Habitat. Sandy beach with seagrass bed, reef flat, intertidal to 45 m.

Distribution. Wide distribution in the Indo-Pacific. Indonesia: Maluku and Lombok.

Remarks. The pale green or light brown color of the chelipeds and ambulatory legs, the whitish cream with dark brown band medially to bright green corneas of the ocular peduncles, and the sea anemones that this species often carries on its gastropod shells are the perfect camouflage in its habitat.

Dardanus guttatus ( Olivier, 1812 )

(Fig. 5B)

Habitat. Sandy beach with seagrass bed and reef flat, intertidal to 3‒4 m.

Distribution. Wide distribution in the Indo-Pacific. Indonesia: Maluku and Lombok.

Remarks. The blue round patch on the merus of the chelipeds and ambulatory legs easily separates D. guttatus from congeners. Gastropods with narrow aperture are very often the preferred shelter of this species.

Dardanus lagopodes ( Forskål, 1775 )

(Fig. 6A)

Habitat. Sandy shore, seagrass bed, and reef flat, intertidal to shallow subtidal.

Distribution. Wide distribution in the Indo-Pacific, from the Red Sea. Indonesia: Lombok, Sulawesi, and Maluku.

Remarks. The color of the specimen in this study is slightly brighter than the specimens from Panglao examined by Malay et al. (2018). However, the shape and ornamentation of the left third pereopod which is flattened, broad, and covered with corneous spines give no doubt of its identity as D. lagopodes.

Figure 6.(A)
Dardanus lagopodes (Forskål, 1775), female, 5.5 mm (MZB Cru 5519). (B) Dardanus scutelatus (H. Milne Edwards, 1848), male, 3.6 mm (MZB Cru 5523). (C) Clibanarius ambonensisRahayu and Forest, 1993 (photo in preservative), male, 5.3 mm (MZB Cru 5524). (D) Clibanarius cruentatus (H. Milne Edwards, 1848), male, 4.9 mm (MZB Cru 5525). (E) Clibanarius demaniBuitendijk, 1937, male, 2.7 mm (MZB Cru 5526). (F) Clibanarius englaucusBall and Haig, 1972, male, 2.2 mm (MZB Cru 5529). Live coloration unless indicated otherwise.

Dardanus megistos (Herbst, 1804)

(Fig. 5E, F)

Habitat. Sandy muddy shore, seagrass bed, and reef flat, intertidal to shallow subtidal.

Distribution. Wide distribution in the Indo-Pacific. Indonesia: Lombok, Sulawesi, and Maluku.

Remarks. This is a very common species in the littoral tropical area of the Indo-West Pacific, and is easily recognized by the bright red color with white spots on the shield, abdomen, chelipeds, and ambulatory legs.

Dardanus scutellatus (H. Milne Edwards, 1848 )

(Fig. 6B)

Habitat. Sandy shore, seagrass bed, and reef flat, intertidal.

Distribution. Indian Ocean to the Philippines. Indonesia: Timor and Maluku.

Remarks. Only one specimen was collected during this study. The specific characters include the elongated left cheliped palm that is covered by short stiff setae, with a denticulate lower margin continued to the fixed finger, and the sculptured lateral face of the left third pereopod. This is not a very common species in Indonesian waters.

Family Diogenidae Ortmann, 1892

Genus Clibanarius Dana, 1852

Clibanarius ambonensis Rahayu and Forest, 1993

(Fig. 6C)

Habitat. The habitat ranges from seagrass, rocky area, and sandy muddy substrate. In Halmahera, it was found near the estuarine area, upper intertidal.

Distribution. Ryu Kyu Island, Japan. Indonesia: Ambon in central Maluku, Timika, Papua, and now in Halmahera, north Maluku.

Color in life. Shield whitish with brown spots. Ocular peduncles bluish-white, dorsal surface with narrow dark brown longitudinal stripe; proximal part brown. Ocular acicles brown with bluish-white spines. Antennal peduncles bluish-white with dark brown longitudinal stripe on dorsal surface of fifth segment. Chelipeds brown with light blue spines; dactyls bluish white with 2 longitudinal brown stripes on dorsal surfaces. Ambulatory legs bluish-white with brown longitudinal stripes on lateral surfaces: dactyls and meri with 3 longitudinal stripes, propodi with 4 longitudinal stripes, carpi with 2 longitudinal stripes (after Rahayu, 2003).

Remarks. A number of species show longitudinal stripes on the ocular peduncles and ambulatory legs, but without stripes on the shield, namely C. antennatusRahayu and Forest, 1993, C. bistriatus Rahayu and Forest, 1993, C. infraspinatus (Hilgendorf, 1869), and C. clibanarius (Herbst, 1791). Clibanarius ambonensis is easily separated by the coloration in life mentioned above. The related species, C. infraspinatus is reddish brown with yellowish-white or orange stripes on the ambulatory legs, and brown stripes on the ocular peduncles (Rahayu, 2022: fig. 1A, B). In C. clibanarius the ambulatory legs are maroon with cream stripes, the ocular peduncle brown with dark brown longitudinal stripe (Komai et al, 2019: fig. 5), and in C. antennatus it is bluish-white with brown stripes on the ambulatory legs, the thin brown stripe in bluish white on the ocular peduncles is interrupted (Rahayu, 2003). Clibanarius bistriatus has orangish cream ambulatory legs with broad greenish brown longitudinal stripe flanked by very narrow dark brown stripe, the ocular peduncles are cream with dark brown longitudinal stripe.

Clibanarius cruentatus (H. Milne Edwards, 1848 )

(Fig. 6D)

Habitat. Shore with a sandy, muddy substrate, gravels, and boulders, intertidal.

Distribution. Wide distribution in the Indo-West Pacific. Indonesia: Lombok, Sulawesi, and Halmahera.

Remarks. This is not a very common species, but when found, it congregates in large numbers under boulders. The bright yellow spots on the dark brown base color of the shield, chelipeds, and ambulatory legs easily separated this species from congeners.

Clibanarius cf. demani Buitendijk, 1937

(Fig. 6E)

Habitat. Sandy muddy substrate, intertidal.

Distribution. Pulau Kumo, Halmahera.

Remarks. The single specimen in this study is tentatively identified as C. demani in having the following characters: antennular peduncles shorter than ocular peduncles, antennal peduncles reaching only half length of ocular peduncles, propodus of pereopods shorter than dactylus. However, the coloration is somewhat different. Rahayu and Wahyudi (2008) gave the color of C. demani as light green speckled with brown on the shield, the ocular peduncles light green with red spot proximally, the ambulatory legs light green with fine brown longitudinal stripes. In the specimen from Halmahera, the shield is greenish cream speckled with dark brown, the ocular peduncles are dark green without any spot, and the pereopods are dark green with very fine light brown longitudinal stripes. Variation in color and color pattern in hermit crabs have been studied largely, and can be regarded as intraspecific or interspecific variation (see Yoshikawa et al, 2020). The different color of the specimen in this study with C. demani warrants further study using molecular analysis to confirm its identity.

Clibanarius englaucus Ball and Haig, 1972

(Fig. 6F)

Habitat. Sandy shore with gravels and boulders, in upper intertidal, very often found in large numbers under boulders.

Distribution. Mayotte Island, the Indian Ocean, the Philippines, and New Guinea. Indonesia: Lombok and Maluku.

Remarks. Described from New Guinea by Ball and Haig (1972) this species is recognizable by the blue and orange color on the dactylus and distal area of the propodus of the second and third pereopods.

Clibanarius eurysternus Hilgendorf, 1879

(Fig. 7A)

Habitat. Sandy mud substrate with gravel and boulders, intertidal to 1 m.

Distribution. Wide distribution in the Indo-west Pacific, from the East coast of Africa to the Philippines, and Vietnam. Indonesia: Sulawesi and Maluku.

Remarks. The species is easily recognized by the dark brown longitudinal stripes on yellow base color of the shield, ocular peduncles, chelipeds, and ambulatory legs. It usually prefers gastropod with a narrow aperture as shelter, therefore the thorax is very often flattened.

Figure 7. (A)
Clibanarius eurysternus Hilgendorf, 1878, male, 3.5 mm (MZB Cru 5530). (B) Clibanarius longitarsus (De Haan, 1849), (photo in preservative), male, 2.7 mm (MZB Cru 5534). (C) Clibanarius merguiensisDe Man, 1888, male, 3.5 mm (MZB Cru 5538). (D) Clibanarius snelliusiBuitendijk, 1937 (photo in preservative), male, 3.3 mm (MZB Cru 5539). (E) Clibanarius striolatusDana, 1852, male, 6.4 mm (MZB Cru 5541). (F) Clibanarius virescens (Krauss, 1843), male, 4.1 mm (MZB Cru 5544). Live coloration unless indicated otherwise.

Clibanarius longitarsus (De Haan, 1849)

(Fig. 7B)

Habitat. Mostly found in a mangrove environment.

Distribution. Wide distribution in the Indo-West Pacific, from the Red Sea across the Indian Ocean, and Malay Archipelago to Japan. In Indonesia: the species was found in almost all islands in Indonesia, from Jawa to Papua.

Color in life. Shield green-brown with darker brown mottling and patches. Ocular peduncle dorsally olive green, ventrally very pale green or cream. Cheliped olive or brown with irregular row of paler blue or blue-green tubercles and spines, many with corneous tip on dactyl and propodus. Ambulatory legs greenish brown or blackish brown with longitudinal bright blue stripe on lateral faces (after Rahayu and Wahyudi, 2008).

Rahayu (2003) recorded color variation of the specimens from Papua, Indonesia, while later (Rahayu, 2022) she provided color pictures of the specimen from Singapore.

Clibanarius merguiensis De Man, 1888

(Fig. 7C)

Habitat. Sandy, rocky shore, seagrass bed, and reef flat, intertidal.

Distribution. West Indian Ocean to Thailand, Vietnam and the Malay Archipelago, and the Philippines and Taiwan. Indonesia: Lombok, Sulawesi and Maluku.

Remarks. In life the species is recognizable by the yellowish orange with tinge of blue broad stripe on the dark brown or black of ambulatory legs. This color fades rapidly in the preservative. Color variation discussed by Rahayu and Forest (1993) were also found in the specimens in this study:

Clibanarius snelliusi Buitendijk, 1937

(Fig. 7D)

Habitat. Sandy shore with gravel and boulders, and rocky substrate, intertidal to 2 m.

Distribution. Vietnam, New Guinea, and Taiwan. Indonesia: Lombok, Flores, and Maluku.

Color in life. Shield reddish purple or reddish brown with some white patches, posterior carapace green-brown. Ocular peduncles brown with white ring at base of cornea. Chelipeds reddish-purple with some white patches and white spines. Ambulatory legs reddish purple with white patches, longitudinal blue stripe along lateral surface (after Rahayu and Wahyudi, 2008).

Remarks. This is not a very common species, but recognizable by its coloration mentioned above.

Clibanarius striolatus Dana, 1852

(Fig. 7E)

Habitat. Mud substrate near mangrove or seagrass bed, intertidal.

Distribution. From the Red Sea, and the Indian Ocean to the Philippines, Taiwan, and Japan. Indonesia: Jawa, Kalimantan, Lombok, Sulawesi, and Maluku.

Remarks. This is a very common species in Indonesian waters. The dark brown stripes on the light green ambulatory legs easily separated this species from the congeners.

Clibanarius virescens ( Krauss, 1843 )

(Fig. 7F)

Habitat. Sandy muddy substrate at seagrass bed and rocky shore, intertidal.

Distribution. East coast of Africa to Thailand, Taiwan, Japan, and Fiji. Indonesia: Kalimantan, Lombok, Sulawesi, and Maluku.

Remarks. The yellow or yellowish cream of dactylus of the second and third pereopods, and the green or dark green of the propodus, carpus and merus are characteristic for this species. Morgan (1985) and Yoshikawa et al. (2020) recorded the color variation of this species.

Genus Diogenes Dana, 1851

Diogenes avarus Heller, 1865

(Fig. 8A)

Habitat. Sandy, muddy and gravel shore, reef flat, intertidal.

Distribution. From East Africa and the Red Sea across the Indian Ocean to Thailand, the Malay Archipelago, Vietnam, the Philippines, and northern Australia. Indonesia: Lombok, Kalimantan, Sulawesi, Maluku, and Papua.

Remarks. This is a very common species on sandy muddy shores. Forest (1956), McLaughlin and Clark (1997), Rahayu and Forest (1995), Rahayu and Komai (2000), Rahayu and Hortle (2002), Siddiqui et al. (2004) have discussed the variability of the following morphological characters: proportion of ocular peduncles against corneas and shield; presence or absence of spine on fourth segment of antennal peduncles; shape of left cheliped; presence or absence of row of spines on dorsal margin of propodus and carpus of second and third pereopods. The color in life of the specimens in this study is different than those described by Morgan (1987a) for the specimens from Australia, and Haig and Ball (1988) for the specimens from Banda, Maluku. In the present specimens the color is as follows: ocular peduncles blue with oblique dark brown band proximally, left cheliped dark brown with dactyl and fixed finger light brown, ambulatory legs dactyl and merus light brown with dark brown or black band proximally, while the propodus and carpus brownish blue distally, dark brown or black proximally (Fig. 8A). The color of the specimens from Australia is as follows: ocular peduncles cream sometimes with blue-green fleck proximally, cheliped cream with variable area of brown especially on propodus ventrally, ambulatory legs cream, propodus brown near mid-length, and scattered brown areas on carpus and merus. Whereas the specimen from Banda has ocular peduncles uniform light brown, or brown with broad, oblique white band medially, carpus and merus of large cheliped solid light brown, chela white; dactylus of ambulatory legs dark brown proximally and white distally; propodus white medially, dark brown proximally and light brown distally; carpus light brown with white band distally; merus with two brown bands.

Considering the variability of morphological and color characters mentioned above, it is possible that more than one species has been identified as D. avarus in different areas of the Indo-West Pacific. It is beyond the scope of this study to resolve the validity of D. avarus across its distribution.

Figure 8. (A)
Diogenes avarusHeller, 1865, male, 3.5 mm (MZB Cru 5533). (B) Diogenes forestiRahayu and Hortle, 2002, 3.4 mm (MZB Cru 5547). (C) Diogenes pallescensWhitelegge, 1897 (photo in preservative), ovigerous female, 1.6 mm (MZB Cru 5550). (D) Diogenes spinicarpus Rahayu and Forest, 1995 (photo in preservative), ovigerous female, 2.2 mm (MZB Cru 5554). (E) Diogenes tumidus Rahayu and Forest, 1995 (photo in preservative), ovigerous female, 2 mm (MZB Cru 5557). (F) Pseudopaguristes monoporus (Morgan, 1987), male, 4.7 mm (MZB Cru 5559). Live coloration unless indicated otherwise.

Diogenes foresti Rahayu and Hortle, 2002

(Fig. 8B)

Habitat. Sandy muddy substrate, 0‒2 m.

Distribution. Indonesian Papua and Halmahera.

Color in life. Shield cream speckled with light brown; ocular peduncles cream, reddish brown ring on base of cornea, mesial face with light brown stripe starting from base of cornea to base of ocular peduncle, dark brown spot on lateral face proximally; ocular acicle and intercalary rostral process greenish brown; cheliped light brown; second and third pereopod light brown with median large dark brown bands, and streak of greenish brown on each propodus, carpus, and merus.

Remarks. This is the first report of the species after its description.

Diogenes pallescens Whitelegge, 1897

(Fig. 8C)

Habitat. Sandy, muddy shore, intertidal.

Distribution. Wide distribution in the Indo-West Pacific. Indonesia: Maluku, Lombok.

Color in life. Shield mottled with light and dark brown, and white; ocular peduncles cream, with brown ring subdistally, reticulated pattern of light brown on dorsal surface. Chelipeds and ambulatory legs cream mottled with dark and light brown; submedian black spot on dorsal margins of propodi, carpi, and meri of pereiopods (after Rahayu, 2022).

Remarks. Diogenes pallescens was recorded from Maluku as D. sereneiForest, 1956, and D. gardineriAlcock, 1905 (Rahayu and Forest, 1995). The two latter taxa were synonymized to D. pallescens by McLaughlin (2002a). Rahayu and Osawa (2012) discussed the morphological variation that was not mentioned by McLaughlin (2002a) including the presence of a row of spines on the ventromedial margin of the ischium of each cheliped and a prominent spine on the dorsal margin of each merus of the second pereopods, telson lacks the median cleft, but has large marginal spines. The specimens in this study agree well with McLaughin (2002a) and Rahayu and Osawa (2012).

Diogenes spinicarpus Rahayu and Forest, 1995

(Fig. 8D)

Habitat. Sandy muddy substrate, intertidal.

Distribution. Singapore. Indonesia: Sulawesi, Maluku, and Papua.

Color in life. Shield cream with light brown speckles on dorsal surface laterally; ocular peduncle cream with median dark brown spot dorsolaterally, light brown blotch mesially, brown ring at base of cornea; corneas dark brown; chelipeds whitish cream, light brown band on carpus medially, light brown marking on merus distally; second and third pereopods dactyl whitish cream, with brown band proximally, propodus, carpus and merus cream with light brown marking, dark brown band medially.

Remarks. Diogenes spinicarpus belongs to the group of Diogenes that have antennal peduncles shorter than the ocular peduncles and the antennal flagella lack paired long setae (Asakura and Godwin, 2006). The characters that separate D. spinicarpus from the closely related species are as follows: ocular peduncles inflated on corneas and more so basally; antennal acicles short and slender only slightly overreach distal margin of fourth antennal peduncular segment; intercalary rostral process long and broadly triangular; cheliped long or stout, but always covered by small spines; and third pereopod with several teeth on dorsal margin of carpus.

Diogenes tumidus Rahayu and Forest, 1995

(Fig. 8E)

Habitat. Sandy, muddy, and rubbles substrate near mangrove, sometimes on seagrass bed; 0‒2m.

Distribution. Singapore and Thailand. Indonesia: Kalimantan, Sulawesi, Maluku, and Papua.

Color in life. Shield mottled with light brown and cream with dark brown spot medially; ocular peduncle creamy white with dark brown streak proximal to corneas to proximal part, corneas dark grey with reddish brown streak at base. Chelipeds generally light brown with dark brown streak on dorsolateral surface of palm and carpus, spines white. P2 and P3 light brown or cream with dark brown streak on dorsolateral margin of dactyls, propodi and carpi (after Rahayu, 2022).

Remarks. This is a very common species in the intertidal. The specific characters include the shape of the right cheliped which has smooth, subcylindrical dactylus, forming large hiatus between fingers, and relatively dense setae on the chelipeds and ambulatory legs.

Genus Pseudopaguristes McLaughlin, 2002 b

Pseudopaguristes monoporus ( Morgan, 1987 b)

(Fig. 8F)

Habitat. Sandy, muddy and gravel shore, reef flat; intertidal to 20 m.

Distribution. Thailand, Philippines, Singapore, and Guam. Indonesia: Lombok and Maluku.

Remarks. Originally described as Paguristes monoporus by Morgan (1987b), this species possesses only a left gonopore in females, the size of the first left pleopod is reduced and the left second pleopod is absent in males, only eight functional gills are present. This species was transferred to PseudopaguristesMcLaughlin, 2002b, by Rahayu (2005). This is a very common species on sandy and muddy shores. The color of the specimens in this study was lighter than those described by Rahayu (2022) for the specimens from Singapore. However, the red longitudinal stripe on the ocular peduncles and the blue color of the antennular peduncles, specific characters for this species, were still noticeable.

Family Paguridae Latreille, 1802

Genus Pagurus Fabricius, 1775

Pagurus fungiformis Komai and Rahayu, 2004

(Fig. 9A)

Habitat. Sandy shore with gravel and boulders, intertidal.

Distribution. Philippines, and Indonesia: Lombok, Sulawesi, and Ambon, and now Halmahera.

Remarks. The color and color pattern of this species resemble that of P. moluccensis, but the two taxa were easily separated by the flattened, closely set, marginally denticulate tubercles on the chelipeds of P. fungiformis. Differences between P. fungiformis and P. moluccensis have been discussed at length by Komai and Rahayu (2004).

Figure 9. (A)
Pagurus fungiformisKomai and Rahayu, 2004, male, 2.9 mm (MZB Cru 5564). (B) Pagurus hirtimanusMiers, 1880, male, 4.7 mm (MZB Cru 5567). (C) Pagurus moluccensisHaig and Ball, 1988 (photo in preservative),1 male, 1.8 mm (MZB Cru 5569). (D) Pagurixus anceps (Forest, 1954) (photo in preservative), male, 1.3 mm (MZB Cru 5570). (E) Pagurixus haigae Komai and Osawa, 2007 (photo in preservative), male, 2 mm (MZB Cru 5571). (F) Trichopagurus macrochela Komai and Osawa, 2005 (photo in preservative), ovigerous female, 1.8 mm (MZB Cru 5572). Live coloration unless indicated otherwise.

Pagurus hirtimanus Miers, 1880

(Fig. 9B)

Habitat. Sand, gravel substrate near seagrass bed, reef flat; intertidal to 3 m.

Distribution. Wide distribution in the Indo-West Pacific. Indonesia: Lesser Sunda Islands, Sulawesi, and Maluku.

Remarks. The specific characters of this species include: short and stout ocular peduncles, moderately dense setae on chelipeds, second and third pereopods; right cheliped with median row of moderately strong spines on dorsal surface of dactylus, flattened fixed finger, covered with tubercles, palm with strong spines medially, and two rows of spines on dorsal surface of the carpus with some capsulate spines on distal margin. The blue or black median band that separated distal blue and proximal white area of the ocular peduncles, and blue antennular peduncles are also characteristic for this species.

Pagurus moluccensis Haig and Ball, 1988

(Fig. 9C)

Habitat. Sandy shore with gravel and boulders, intertidal to 5 m.

Distribution. Philippines, and Indonesia: Lombok and Maluku.

Color in life. Shield with black markings on white background. Ocular peduncles with broad, subproximal dark band. Antennular peduncles white, penultimate segment with broad, submedian dark band. Antennal peduncles with alternating black and white bands. Chelipeds with complex black pattern on white background; second and third pereopods with black, irregular-edged bands on white background (after Haig and Ball, 1988).

In preservative the black marking became dark brown.

Genus Pagurixus Melin, 1939

Pagurixus anceps ( Forest, 1954 )

(Fig. 9D)

Habitat. Sandy substrate on a rocky shore, intertidal.

Distribution. Wide distribution in the Indo-West Pacific, from Red Sea, Madagascar across the Indian Ocean to Thailand, Philippines, Japan, New Caledonia, and Tuamotu, French Polynesia. Indonesia: Seram and Halmahera in Maluku.

Color in life. Shield generally whitish, with some brown spots anteriorly. Ocular peduncle also whitish, with two brown spots proximal to base of cornea and two spots basally. Antennular peduncle generally transparent; ultimate segment with brown band in proximal 0.3; penultimate segment with brown spot basally; dorsal flagellum pale yellowish-brown. Antennal peduncle also with brown spots on second segment; fifth segment with brown lateral and mesial margins; distal half of antennal acicle brown; antennal flagellum pale brown, without conspicuous markings. Chelipeds generally white, occasionally with tinge of brown on chelae and carpi; meri occasionally with subdistal brown ring. Ambulatory legs white with brown markings; dactyli each with brown ring occupying proximal 0.2-0.3; propodi each with two brown rings, distal one distal to mid-length and proximal one at proximal 0.2, rings occasionally combined with longitudinal bar on lateral surface (rings occasionally broken into dorsal and ventral spots); carpi each with two or three brown spots laterally; meri each with brown subdistal ring (occasionally broken into two spots) and one proximal brown spot (after Komai, 2010).

Remarks. The single male in this study agrees well with the redescription of the species by Komai (2010) on the following characters: ocular peduncles stout, right cheliped strong, carpus without dorsomesial spines, and merus without conspicuous spines on ventrolateral margin, third pereopod relatively stout with only five dactylar spines ventrally.

Although widely distributed in the Indo-West Pacific with a good number of specimens, the specimen in this study is the second record in Indonesian waters after Komai’s (2010) record from Seram Island. The small-size individuals (less than 2.0 mm; see Komai, 2010) are very often overlooked in the field.

Pagurixus haigae Komai and Osawa, 2007

(Fig. 9E)

Habitat. Sandy substrate on a rocky shore, intertidal, under large boulders.

Distribution. Widely distributed in the Indo-West Pacific, from western Indian Ocean to Thailand, Malaysia, Philippine, Japan, Guam and New Caledonia; Indonesia: Maluku.

Color in life. Shield with black markings on white background. Ocular peduncles with broad, subproximal dark band. Antennular peduncles white, penultimate segment with broad, submedian dark band; antennal peduncles with alternating black and white bands. Chelipeds with complex black pattern on white background; second and third pereopods with black, irregular-edged bands on white background (after Komai and Osawa, 2007).

Remarks. Two specimens were collected in this study. In the preservative the species still shows the distinctive longitudinal brownish-red longitudinal stripes on pereopods and chelipeds.

Genus Trichopagurus de Saint Laurent, 1968

Trichopagurus macrochela Komai and Osawa, 2005

(Fig.9F)

Habitat. Sandy substrate on a rocky shore, among dead coral, intertidal.

Distribution. Philippines and Ryukyu, Amami Island and Yap Island, Japan. This is a new record in Indonesian waters.

Color in life. Carapace, antennae, chelipeds and ambulatory legs generally pale reddish brown, without conspicuous markings; setae at base of cornea accentuated with dark reddish brown (after Komai and Osawa, 2005).

Remarks. Although only one ovigerous female specimen was collected in this study, the morphological characters agree well with T. macrochela: 11 pairs of biserial gills, carpus of right cheliped flattened, dorsomesial margin with row of strong spines, female with left gonopore only, fifth coxae asymmetrical, left larger than right. Slight differences with Komai and Osawa’s (2005) descriptions is that in the material in this study the fifth coxae both with numerous long setae on the ventral surface.

DISCUSSION

This is the first time that the hermit crabs are reported from the northeast coast of Halmahera Island, northern part of Maluku in Indonesia, therefore all species are new records for the area. Considering that the hermit crabs in this study were collected by hand intertidally, from six different localities, the 34 species obtained were relatively numerous. For comparison, during Alpha Helix expedition 1975, in central part of Maluku (Ambon, Seram and Banda Islands), Haig and Ball (1988) reported 44 species of hermit crabs, that were collected by various methods, including scuba diving, free diving, and intertidal collection. Seventeen species reported by Haig and Ball (1988) were also found in this study, namely Coenobita rugosus, Ciliopagurus strigatus (as Trizopagurus strigatus), Calcinus gaimardii, C. laevimanus, C. latens, Clibanarius englaucus, C. eurysternus, C. longitarsus, C. virescens, Dardanus lagopodes, D. megistos, D. scutellatus, Diogenes avarus, Diogenes pallescens (as D. serenei), Pseudopaguristes monoporus (as Paguristes monoporus), Pagurus hirtimanus, and P. moluccensis. Given the same collecting methods, hermit crabs on the northern coast of Halmahera may be as many as in Central Maluku.

ACKNOWLEDGEMENTS

We thank Egenius Resusun and Welhelmus Sahuleka for helping in the field. Comments and suggestions from the reviewers helped to improve the manuscript

REFERENCES

• Alcock A 1905. Catalogue of the Indian decapod Crustacea in the collections of the Indian Museum Part 2. Anomura. Fasc. 1. Pagurides Indian Museum, Calcutta, xi + 197 pp. https://doi.org/10.5962/bhl.title.10323
  » https://doi.org/10.5962/bhl.title.10323
• Asakura A and Godwin S 2006. Diogenes patae n. sp., a new species of hermit crab (Crustacea, Decapoda, Anomura, Diogenidae) from American Samoa. Zoosystema, 28 (2): 457‒463. https://doi.org/10.5281/zenodo.5402845
  » https://doi.org/10.5281/zenodo.5402845
• Ball EE and Haig J 1972. Hermit crabs from eastern New Guinea. Pacific Science, 26: 87‒107.
• Buitendijk AM 1937. Biological results of the Snellius Expedition. IV. The Paguridea of the Snellius Expedition. Temminckia, 2: 251-280.
• Dana JD 1851. Conspectus Crustaceorum quae in orbis terrarium circumnavigatione, Carolo Wilkes e classe Reipublicae Foederatae duce, lexit et descripcit. Proceeding of the Academy of Natural Sciences, Philadelphia, 5: 267-272.
• Dana JD 1852. Crustacea, part I. United States Exploring Expedition, during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U.S.N., Vol. 13. C. Sherman, Philadelphia (Reprinted Antiquariaat Junk, Lochem, Netherlands, 1972).
• De Haan W 1833-1850. Crustacea. In: Siebold PF von (ed.) Fauna Japonica sive Descriptio Animalium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui Summum in India Batava Imperium Tenent, Suscepto, Annis 1823-1830 Collegit, Notis, Observationibus et Adumbrationibus Illustravit. Lugduni-Batavorum, Leiden. 243p.
• De Man JG 1888. Report on the Podophthalmous Crustacea of the Mergui Archipelago, collected for the Trustees of theIndian Museum, Calcutta , by Dr. John Anderson, F.R.S., Superintendent of the Museum, parts IV and V. Journal of the Linnean Society, London, 22: 177-240, 241-305.
• Fabricius JC 1775. Systema Entomologiae, sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus , observationibus. Officina Libraria Kortii. 832 pp.
• Forest J 1952. Contributions à la révision des crustacés Paguridae. I. Le genre Trizopagurus Mémoires du Muséum national d’Histoire naturelle, serie A, Zoologie, 5 (1): 1‒40.
• Forest J 1954. Crustacés Décapodes Marcheurs des îles de Tahiti et des Tuamotu. - I. Paguridea. Cont. Bulletin du Muséum national d’Histoire naturelle, (1) 25: 71‒79.
• Forest J 1956. Les Pagures du Viet-Nam. I. Le genre Diogenes Dana. Bulletin du Muséum national d’Histoire naturelle, Series 2, 28: 524-532.
• Forest J 1995. Crustacea Decapoda Anomura: révision du genre Trizopagurus Forest, 1952 (Diogenidae) avec l’établissement de deux genres nouveaux. In: Crosnier, A (Ed.), Résultats des campagnes MUSORSTOM, volume 13. Mémoires du Muséum national d’Histoire naturelle, 163: 9‒149.
• Forskål P 1775. Descriptiones Animalium, Avium, Amphibiorum, Piscium, Insectorum, vermium, quae in itinere oriental observ avit. Post mortem auctoris edidit Carsten Niebuhr. Hauniæ, pp. 1-19, 1-164.
• Fraajie RHB, Van Bakel BWM and Jagt JWM 2017. A new paguroid from the type Maastrichitian (upper Crestaceous, the Netherland) and erection of a new family. Earth Science Bulletin, 188: 17. https://doi.org/10.1051/bsgf/2017185
  » https://doi.org/10.1051/bsgf/2017185
• Haig J and Ball EE 1988. Hermit crabs from northern Australian and eastern Indonesian waters (Crustacea Decapoda: Anomura: Paguroidea) collected during the 1975 Alpha Helix Expedition. Records of the Australian Museum, 40: 151-196.
• Heller C 1865. Crustaceen. In: Reise der Österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodors B. von Wüllerstorf-Urbair. Zoologischer Theil, 2(3): 1‒280, pls. 1‒25. Wien, Kaiserlich-königlichen Hof-und Staatsdruckerei.
• Herbst JFW 1791-1804. Versuch einer Naturgeschichte der Krabben und rebse nebst einer systematischen Beschreibung ihrer verschiedenen Arten. 2: 1-226. Berlin, Bei Gottlieb August Lange. https://doi.org/10.5962/bhl.title.64679
  » https://doi.org/10.5962/bhl.title.64679
• Hilgendorf F 1869. Crustaceen. In: van der Decken CC (Ed.), Reisen in Ost-Afrika in dem Jahren 1859-1865, 3 (1): p. 69-116. Leipzig und Heidelberg, C.F. Winter’sche Verlagshandlung,
• Komai T 2004. Calcinus areolatus Rahayu and Forest, an invalid taxon founded on juvenile or young stages of Calcinus morgani Rahayu and Forest (Crustacea: Decapoda: Anomura: Diogenidae). Biology Magazine of Okinawa, 42: 25-38.
• Komai T 2010. New species and new records of the hermit crab genus Pagurixus Melin, 1939 (Crustacea: Decapoda: Anomura: Paguridae) from the Indo-West Pacific. Journal of Natural History, 44: 21, 1269-1342. https://doi.org/10.1080/00222931003624812
  » https://doi.org/10.1080/00222931003624812
• Komai T and Osawa M 2005. A new species of Trichopagurus de Saint Laurent (Crustacea:Decapoda: Anomura: Paguridae) from the Ryukyu and Yap Islands, and redescription of T. trichophthalmus (Forest). Zootaxa, 801: 1-20. http://dx.doi.org/10.11646/zootaxa.5419.1.5
  » http://dx.doi.org/10.11646/zootaxa.5419.1.5
• Komai T and Osawa M 2007. A new species of the hermit crab genus Pagurixus Melin (Crustacea: Decapoda: Anomura: Paguridae) from the Indo-West Pacific. Raffles Bulletin of Zoology, 55: 97‒105. https://doi.org/10.5281/zenodo.5354241
  » https://doi.org/10.5281/zenodo.5354241
• Komai T and Rahayu DL 2004. Redescription of Pagurus moluccensis Haig and Ball, 1988, and description of a new species of Pagurus from Indonesia (Crustacea: Decapoda: Anomura: Paguridae). Raffles Bulletin of Zoology, 52: 183-200. https://doi.org/10.5281/zenodo.4618844
  » https://doi.org/10.5281/zenodo.4618844
• Komai T, Reshmi R, Rahayu DL and Kumar AB 2019. Redescription of the little known hermit crab, Clibanarius clibanarius (Herbst, 1791) (Decapoda: Anomura: Diogenidae), based on material from southern India and type material. Zootaxa, 4555 (3): 372‒384.https://doi.org/10.11646/zootaxa.4555.3.6
  » https://doi.org/10.11646/zootaxa.4555.3.6
• Krauss F 1843. Die Südafrikanischen Crustaceen. Eine Zusammenstellung aller bekannten Malacostraca. Bemerkungen über deren Lebensweise und geographische Verbreitung, nebst Beschreibung und Abbildung mehrerer neuen Arten. Stuttgart, E. Schweizerbart’sche Verlagsbuchhandlung, 68p.
• Latreille PA 1802. Histoire naturelle, générale at particulière, des Crustacés et des Insects, Vol. 3. Paris, F. Dufart, 467p.
• Latreille PA 1829. Crustacé, Arachnides et partie des Insectes. In: Cuvier G. Le règne animal distribute d’après son organization, pour server de base à l’histoire naturelle des animaux et d’introduction à l’anatomie compare. Deterville, Paris. Pp.1‒39.
• Malay MC, Rahayu DL and Chan T-Y 2018. Hermit crabs of the genera Calcinus Dana, Clibanarius Dana, and Dardanus Paul’son from the PANGLAO 2004 Expedition, with description of a new species and a checklist of the hermit crabs of the Philippines (Crustacea: Anomura: Paguroidea). Raffles Bulletin of Zoology, 66: 23‒65. https://doi.org/10.5281/zenodo.4503028
  » https://doi.org/10.5281/zenodo.4503028
• McLaughlin PA 2002a. Diogenes pallescens Whitelegge, D. gardineri Alcock, D. serenei Forest (Decapoda, Anomura, Diogenidae): morphological variant or distinct species? Raffles Bulletin of Zoology, 50: 81-94.
• McLaughlin PA 2002b. Pseudopaguristes, a new and aberrant genus of hermit crabs (Anomura: Paguroidea: Diogenidae). Micronesica, 34(2): 185‒199.
• McLaughlin PA and Clark P 1997. A review of the Diogenes (Crustacea, Paguroidea) hermit crabs collected by Bedford and Lanchester from Singapore, and from the ‘Skeat’ Expedition to the Malay Peninsula, with a description of a new species and notes on Diogenes intermedius De Man, 1892. Bulletin of the Natural History Museum, London (Zoology), 63: 33‒49. https://biostor.org/reference/113711
  » https://biostor.org/reference/113711
• McLaughlin PA, Rahayu DL, Komai T and Chan T-Y 2007. A catalog of the hermit crabs (Paguroidea) of Taiwan. Keelung, National Taiwan University, 366p.
• Melin G 1939. Paguriden und Galatheiden von Prof. Dr. Sixten Bocks Expedi tion nach den Bonin-Inseln 1914. Kongliga Svenska Vetenskapsakademiens Handlingar (3)18(2): 1-119.
• Miers EJ 1880. On a collection of crustacea from the Malaysian region Part III. Crustacea Anomura and Macrura (except Penaeidea). Annals and Magazine of Natural History, (5) 5: 370‒384.
• Milne-Edwards H 1836. Observations zoologiques sur les Pagures et description d’un nouveau genre de la tribu des Paguriens. Annales des Sciences Naturelle Zoologie, Paris, 6: 257-288.
• Milne-Edwards H 1837. Histoire Naturelle des Crustacés, Comprenant l´Anatomie, la Physiologie et la Classification de ces Animaux. Vol. 2. Paris, Librairie Encyclopédique Roret, 532p.
• Milne-Edwards H 1848. Note sur quelques nouvelles espèces du genre Pagure. Annales des Sciences Naturelles Zoologie, Paris, (3)10: 59-64.
• Morgan GJ 1985. Variation in a key character in a population of Clibanarius virescens (Krauss, 1843) (Decapoda, Diogenidae). Crustaceana, 55: 317‒318.
• Morgan GJ 1987a. Hermit crabs (Decapoda, Anomura: Coenobitidae, Diogenidae, Paguridae) of Darwin and Port Essington, Northern Australia. The Beagle, Records of the Nortern Territory Museum Arts and and Science, 4 (1): 165‒186. https://doi.org/10.5962/p.271277
  » https://doi.org/10.5962/p.271277
• Morgan GJ 1987b. A new and aberrant species of Paguristes (Anomura: Diogenidae) from northern Australia. Records of the Western Australian Museum, 13 (3): 379-386.
• Morgan GJ 1991. A review of the hermit crab genus Calcinus Dana (Crustacea: Decapoda: Diogenidae) from Australia, with descriptions of two new species. Invertebrate Taxononomy, 5: 869‒913. https://doi.org/10.1071/IT9910869
  » https://doi.org/10.1071/IT9910869
• Nakasone Y 1988. Land hermit crabs from the Ryukyus, Japan, with a description of a new species from the Philippines (Crustacea, Decapoda, Coenobitidae). Zoological Science, 5: 165-178.
• Olivier GA 1812. Pagure. Pagurus p. 631‒647. Encyclopedic methodique. Histoire naturelle, Insectes, Vol. 8. Paris, H. Agasse.
• Ortmann A 1892. Die Decapoden-Krebse des Strassburger Museum, mit beson derer Berücksichtigung der von Herrn Dr. Doederlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strassburger Museum aufbe wahrten Formen. IV. Die Abtheilungen Galatheidea und Paguridea. Zoologishen Jahrbüchern. Abtheilung für Systematik, Geographie und Biologie der Thiere, 6: 241-326.
• Paul’son O 1875. Izsledovaniya rakoobraznykh krasnago morya s zametkami otnositel’no rakoobraznykh drugikh morei. Chast’ 1. Podophthalmata i Edriophthalmata (Cumacea), i-xiv + 144 pp. S.V. Kul’zhenko, Kiev. [Studies on Crustacea of the Red Sea with notes regarding other seas. Podophthalmata and Edriophthalmata (Cumacea).] Translation, Israel Program for Scientific Translations, 1961, National Science Foundation and Smithsonian Institution.
• Rahayu DL 2003. Hermit crab species of the genus Clibanarius (Crustacea: Decapoda: Diogenidae) from mangrove habitats in Papua, Indonesia, with description of a new species. Memoires of Museum Victoria, 60(1): 99‒104.
• Rahayu DL 2005. Addition to the Indonesian fauna of the hermit crab genus Pseudopaguristes McLaughlin and a further division of the genus Paguristes Dana (Crustacea: Decapoda: Paguroidea: Diogenidae). Zootaxa, 831: 1‒42. https://doi.org/10.11646/zootaxa.831.1.1
  » https://doi.org/10.11646/zootaxa.831.1.1
• Rahayu DL 2021. Hermit crabs of the family Diogenidae (Crustacea: Decapoda: Anomura) collected during the South Java Deep-Sea 2018 biodiversity cruise in Indonesia, with description of two new species. Raffles Bulletin of Zoology, Suppl. 36: 162‒180. https://doi.org/10.26107/RBZ-2021-0036D
  » https://doi.org/10.26107/RBZ-2021-0036D
• Rahayu DL 2022. Hermit crabs of Singapore (Crustacea: Decapoda: Anomura: Diogenidae, Paguridae), with description of two new species. Raffles Bulletin of Zoology, 70: 329-363. https://doi.org/10.26107/RBZ-2022-0017
  » https://doi.org/10.26107/RBZ-2022-0017
• Rahayu DL and Forest J 1993. Le genre Clibanarius (Crustacea, Decapoda, Diogenidae) en Indonésie, avec la description de six espèces nouvelles. Bulletin du Muséum national d’Histoire naturelle, Paris, 14(A): 745-779.
• Rahayu DL and Forest J 1995. Le genre Diogenes (Crustacea, Decapoda, Diogenidae) en Indonésie, avec la description de six espèces nouvelles. Bulletin Muséum National d’Histoire Naturelle, Paris, 16 (2‒4): 383‒415.
• Rahayu DL and Forest J 1999. Sur le statut de Calcinus gaimardii (H. Milne Edwards, 1848) (Decapoda, Anomura, Diogenidae) et description de deux espèces nouvelles apparentées. Zoosystema, 21(3): 461-472.
• Rahayu DL and Hortle KG 2002. The genus Diogenes (Decapoda, Anomura, Diogenidae) from Irian Jaya, Indonesia with description of a new species. Crustaceana, 75 (3‒4): 609‒619.
• Rahayu DL and Komai T 2000. Shallow-water hermit crabs (Crustacea: Decapoda: Anomura: Diogenidae and Paguridae) of Phuket, Thailand. Phuket Marine Biology Center Research Bulletin, 63: 21‒44.
• Rahayu DL and Osawa M 2012. Hermit crabs (Crustacea: Decapoda: Diogenidae) from the KUMEJIMA 2009 Expedition, Japan. In: Naruse T; Chan T-Y ; Tan HH; Ahyong ST and Reimer JD , Scientific Results of the Marine Biodiversity Expedition ‒ KUMEJIMA 2009. Zootaxa, 3367: 176‒190. https://doi.org/10.11646/zootaxa.3367.1.17
  » https://doi.org/10.11646/zootaxa.3367.1.17
• Rahayu DL and Pratiwi R 2022. Diogenes matabiru, a new species of hermit crab from Lombok Island, Indonesia (Crustacea, Decapoda, Anomura, Diogenidae). Zootaxa, 5093(4): 493-500. https://doi.org/10.11646/zootaxa.5093.4.7
  » https://doi.org/10.11646/zootaxa.5093.4.7
• Rahayu DL and Wahyudi AJ 2008. Common littoral hermit crabs of Indonesia. Editors: Susetiono Y, Shirayama Y, Asakura A, Kyoto, Kyoto University Press. 93p.
• Randall JW 1840. Catalogue of the Crustacea brought by Thomas Nuttall and J.K. Townsend, from the west coast of North America and the Sandwich Islands, with descriptions of such species as are apparently new, among which are included several species of different localities, previously existing in the collection of the Academy. Journal of the Academy of Natural Sciences of Philadelphia, 8: 106-147.
• Saint Laurent de M 1968. Révision des genres Catapaguroides et Cestopagurus et description de quatre genres nouveaux. I. Catapaguroides A. Milne Edwards et Bouvier et Decaphyllus nov. gen. (Crustacés Décapodes Paguridae). Bulletin du Muséum National d’Histoire Naturelle (2)39(5)(1967): 923-954.
• Siddiqui FA, Kazmi OB and McLaughlin PA 2004. Review of the Pakistani species of Diogenes Dana, 1951 (Decapoda Anomura Paguroidea Diogenidae). Tropical Zoology, 17: 155‒200.
• Whitelegge T 1897. The fauna of Funafuti [6] The Crustacea. Australian Museum Memoirs, 3: 7‒151.
• Yoshikawa A, Ikeo K, Imoto J, Jaingam W, Putri LSE, Mardiansyah, -, Nakano T, Shimomura M and Asakura A 2020. Colour variation of the intertidal hermit crab Clibanarius virescens considering growth stage, geographic area in the Indo-West Pacific Ocean, and molecular phylogeny. Journal of the Biological Association of the United Kingdom, 100(7): 1107‒1121. https://doi.org/10.1017/S002531542000106X
  » https://doi.org/10.1017/S002531542000106X

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ADDITIONAL INFORMATION AND DECLARATIONS

• Consent for publication

  All authors declare that they have reviewed the content of the manuscript and gave their consent to submit the document
• Funding and grant disclosures This research was funded by the DIPA of the Research Organization Life Science and Environmental SK Number 14/III/HK/2021 and DIPA Deputy for Facilitation of Research and Innovation, National Research and Innovation Agency (BRIN) Fiscal Year 2022

• Study association

  This study was developed in collaboration with Pusat Riset Biosistematika dan Evolusi, Pusat Riset Bioindustri Laut dan Perairan Darat, Direktorat Pengelolaan Koleksi Ilmiah at Badan Riset dan Inovasi Nasional (BRIN) and Universitas Hein Namotemo, Maluku Utara, Indonesia

• Study permits

  Field collection and transportation of specimens were made under the permit from Pusat Riset Bioindustri Laut dan Perairan Darat-BRIN number B-27469/III.4.8/KP.11.00/6/2022

• Data availability

  All study data are included in the article and specimen collections are deposited within Museum Zoologicum Bogoriense (MZB) and available on request to the corresponding author.

Edited by

Associate Editor

Kareen Schnabel

Editor-in-chief

Christopher Tudge

Data availability

All study data are included in the article and specimen collections are deposited within Museum Zoologicum Bogoriense (MZB) and available on request to the corresponding author.

Publication Dates

• Publication in this collection
  29 July 2024
• Date of issue
  2024

History

• Received
  07 Jan 2023
• Accepted
  02 Sept 2023