Integrative Taxonomy Uncovers Four New Species and One New Record of Land Hermit Crabs Coenobita Latreille, 1829 (Crustacea: Decapoda: Anomura: Coenobitidae) from Indonesia

© 2025 Academia Sinica, Taiwan Open Access Integrative Taxonomy Uncovers Four New Species and One New Record of Land Hermit Crabs Coenobita Latreille, 1829 (Crustacea: Decapoda: Anomura: Coenobitidae) from Indonesia Hsi-Te Shih1,§,* , Dwi Listyo Rahayu2,§, and Félix Adhi Pramono3 1Department of Life Science and Global Change Biology Research Center, National Chung Hsing University, Taichung 402, Taiwan. *Correspondence: E-mail: [email protected] (Shih) 2Research Center for Marine and Land Bioindustry, National Research and Innovation Agency, Dusun Teluk Kodek, Pemenang, Lombok Utara 83756, NTB, Indonesia. E-mail: [email protected] (Rahayu) 3Jalan Kacang Polong I #29, Bojong Indah, Rawabuaya - Cengkareng, Jakarta Barat 11740, Indonesia. E-mail: [email protected] (Pramono) urn:lsid:zoobank.org:pub:D2750620-CE61-48C7-9333-D2C43EAB5067 §HTS and DLR contributed equally to this paper. Received 26 July 2024 / Accepted 24 February 2025 / Published -- 2025 Communicated by Benny K.K. Chan In this study, four new species of land hermit crabs (Crustacea: Decapoda: Anomura: Coenobitidae: Coenobita Latreille, 1829) are described from Indonesia. These descriptions are based on evidence from morphological differences as well as mitochondrial 16S rDNA and cytochrome c oxidase subunit I data. Among the new species, Coenobita moluccensis n. sp. (from Aru Island, Maluku), C. patsyae n. sp. (from Central Sulawesi and Southeast Sulawesi), and C. celebensis n. sp. (from Central Sulawesi) form a major clade on the phylogenetic tree, exhibiting similarities in morphology, particularly in the male sexual tubes— a character they share with their closely related counterpart, C. lila Rahayu, Shih & Ng, 2016. Although the four species are similar in overall morphology, they can be distinguished by differences in the left third pereopod, the second article of the antennal peduncles, granulation on the pereopods, as well as their live coloration. Coenobita granularis n. sp., found in Central Sulawesi, shares morphological similarities with the genetically closely related C. pseudorugosus Nakasone, 1988, particularly in the male sexual tubes. However, the two species can be distinguished by differences in the morphology of the male sexual tubes, the presence of tubercles on the left cheliped, the left third pereopod, as well as their live coloration. Additionally, a newly recorded species, C. variabilis McCulloch, 1909, has been confirmed in West Papua. This study brings the total number of Coenobita species known from Indonesia to 13. Key words: Coenobita moluccensis, C. patsyae, C. celebensis, C. variabilis, New species, Newly recorded species, Morphology, 16S rDNA, Cytochrome c oxidase subunit I (COI), Phylogeny Citation: Shih HT, Rahayu DL, Pramono FA. 2025. Integrative taxonomy uncovers four new species and one new record of land hermit crabs Coenobita Latreille, 1829 (Crustacea: Decapoda: Anomura: Coenobitidae) from Indonesia. Zool Stud 64:11. doi:10.6620/ZS.2025.64-11. BACKGROUND The genus Coenobita contains 17 species, distributed throughout the tropical and subtropical zones of the world, including the Indo-West Pacific (IWP) and Atlantic-East Pacific (AEP) regions (Hartnoll 1988; McLaughlin et al. 2010; Rahayu et al. 2016; Shih et al. 2023a). In the Coral Triangle region (approximately corresponding to the ranges of the Malay Archipelago or the Indo-Australian Archipelago [IAA], depending Zoological Studies 64:11 (2025) doi:10.6620/ZS.2025.64-11 1 © 2025 Academia Sinica, Taiwan on the definition; Hoeksema 2007; Lohman et al. 2011), this genus exhibits the highest species diversity, with up to eight species recorded: C. brevimanus Dana, 1852, C. cavipes Stimpson, 1858, C. lila Rahayu, Shih & Ng, 2016, C. longitarsis De Man, 1902, C. perlatus H. Milne Edwards, 1837, C. pseudorugosus Nakasone, 1988, C. rugosus H. Milne Edwards, 1837, and C. violascens Heller, 1862 (Nakasone 1988; Shih 2012 2020; Rahayu et al. 2016; Shih et al. 2023a). This high species diversity is consistent with patterns seen in other coastal organisms in the Coral Triangle, e.g., corals, shrimps, gastropods, and reef fishes (De Grave 2001; Hoeksema 2007; Bellwood and Meyer 2009; Veron et al. 2009). Indonesia is a biodiversity hotspot, boasting the highest marine species diversity among countries within the Coral Triangle, and many marine species remain undescribed (Veron et al. 2009; Siallagan et al. 2023). Currently, all eight species of Coenobita mentioned above have been recorded in Indonesia (Wáng 2006; Rahayu et al. 2016; Shih et al. 2023a). In recent years, one of the authors (F.A. Pramono) collected specimens of Coenobita from the Indonesian coast and observed some individuals displaying distinctive coloration and morphology that warranted further study to clarify their species identity. After a detailed examination of their morphology and analyses of molecular data from cytochrome c oxidase subunit I (COI), four new species and one newly recorded species from Indonesia have been confirmed. Three of the new species belong to the group lacking a stridulatory ridge, while one new species possesses a stridulatory ridge on the outer palm of the left chela. These four species differ from their congeners in several morphological characters, including the left cheliped, the left third pereopods, the second article of the antennal peduncles, the male sexual tubes, and the live coloration. The new record is C. variabilis McCulloch, 1909, which was originally thought to be endemic to Australia. MATERIALS AND METHODS Specimens examined or sequenced were deposited in the Museum Zoologi Bogor (MZB), National Research and Innovation Agency (= Badan Riset dan Inovasi Nasional, BRIN), Cibinong, Indonesia; the Zoological Collections of the Department of Life Science, National Chung Hsing University (NCHUZOOL), Taichung, Taiwan; the Florida Museum of Natural History, University of Florida, Florida, USA (UF); and the Zoological Reference Collection (ZRC) of the Lee Kong Chian Natural History, National University of Singapore, Singapore. Morphological characters were illustrated with the aid of a drawing tube attached to a stereomicroscope. Descriptive terminology followed that of McLaughlin et al. (2007) and Rahayu et al. (2016). The abbreviations P2, P3, P4, and P5 refer to the second, third, fourth, and fifth pereopods, respectively. Specimen size is indicated by shield length (SL) in mm, measured from the tip of the rostrum or midpoint of the rostral lobe to the midpoint of the posterior margin of the shield. The length of the ocular peduncles was measured along the left ultimate peduncular segment, including the cornea, along its lateral surface; and corneal diameter represents the maximum width of the cornea measured on the dorsal surface. Since C. moluccensis, C. patsyae, and C. celebensis are similar in morphology, a full description is provided only for C. moluccensis. However, a Diagnosis section is included for each species (see the example in Shih et al. 2023c). Genomic DNA was isolated from gill or muscle tissue using kits (see Shih et al. 2016 for details). A region of approximately 545 basepairs (bp) from the 5'-end of the mitochondrial large ribosomal subunit (16S rDNA) gene was selected for amplification with polymerase chain reaction (PCR) using the primers 1471, 1472 (Crandall and Fitzpatrick 1996), 16L29, 16H10, and 16H11 (Schubart 2009), as well as the newly designed primers, 16L29B (5'-YGCCTGTTTAYCAAAAACAT-3') and 16H10B (5'-AATCCTTTCGTACTARR-3'). A portion of the COI gene was amplified using PCR with the primers LCO1490, HCO2198 (Folmer et al. 1994), LCOB, HCOex3 (Shih et al. 2022), LCOC, HCOex4 (Shih et al. 2023a), HCOex0 (Shih et al. 2023b), LCOex2, and LCOex3 (Shih et al. 2023d), as well as the newly designed primer, HCOex1 (5'-GCYTCTTTYTTDCCMGACTC-3'). PCR conditions for the above primers involved 40 cycles of denaturation for 50 s at 94°C, annealing for 70 s at 45– 47°C, and extension for 60 s at 72°C, followed by a final extension for 10 min at 72°C. Sequences were obtained by automated Sanger sequencing (Applied Biosystems 3730, USA). Sequences of the different haplotypes were deposited in NCBI GenBank (accession numbers are shown in Table 1). Additional COI sequences published in Rahayu et al. (2016) and Shih et al. (2023a) were also included (Table 1). Sequences were aligned using the MUSCLE function in MEGA (vers. 11, Tamura et al. 2021). For the combined 16S and COI dataset, the bestfitting models for sequence evolution of individual datasets were determined by PartitionFinder (vers. 2.1.1, Lanfear et al. 2017), using the Bayesian information criterion (BIC). The best model, HKY+I+G, was page 2 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan subsequently applied for maximum likelihood (ML) and Bayesian inference (BI) analyses. The ML analysis was conducted in IQ-TREE (vers. 2.2.0, Minh et al. 2020) with the best models and 30,000 ultrafast bootstrap replicates (Hoang et al. 2017). The BI was performed with MrBayes (vers. 3.2.6, Ronquist et al. 2012). The search was run with four chains for 10 million generations and four independent runs, with trees sampled every 1000 generations. The convergence of chains was determined by the average standard deviation of split frequency values below the recommended 0.01 (Ronquist et al. 2020), and the first 3500 trees were discarded as the burnin. A TCS haplotype network of the 16S+COI haplotypes was generated using the program PopART (vers. 1.7, Leigh and Bryant 2015). For barcoding COI, basepair (bp) differences and pairwise estimates of Kimura 2-parameter (K2P) distances (Kimura 1980) for genetic diversities between COI haplotypes were calculated with MEGA. Comparative material: Coenobita cavipes Stimpson, 1858: 1 ♂ (12.4 mm) (ZRC 2013.0267), Manado, Sulawesi, Coll. N.K. Ng, 25 Sep. 2003; 1 ♂ (11.3 mm) (MZB Cru 5724), Ajkwa, Papua, 29 September 2021; 1 ♂ (20.0 mm), Malalayang, Manado, Sulawesi, coll. N.K. Ng, 14 Apr. 2003. C. lila Rahayu, Shih & Ng, 2016: 1 paratype ♂ (10.7 mm) (NCHUZOOL 13624); 1 ♂ (11.3 mm) (NCHUZOOL 15284), St John’s Island, coll. H-T Shih et al., 10 Feb. 2014; 10 ♂♂ (3.8‒11.3 mm), 1 ♀ (4.6 mm), 1 ovig. ♀ (9.0 mm, St John’s Island, Singapore, 10 February 2014; 10 ♂♂ (3.8–11.3 mm), 1 ♀ (4.6 mm), 1 ovig. ♀ (9.0 mm) (ZRC 2013.1782) St 78, St John’s Island, 25 May 2013. C. perlatus H. Milne Edwards, 1837: 2 ♂♂ (10.3, 10.9 mm) (NCHUZOOL 17213), 1 ♀ (13.5 mm) (NCHUZOOL 17230), Pago Bay, Guam, 4 Aug. 2001. C. pseudorugosus Nakasone, 1988: Central Sulawesi, Indonesia: 2 ♂♂ (13.0, 14.2 mm) (NCHUZOOL 17199), Oct. 2022; 6 ♂♂ (8.0–13.8 mm) (NCHUZOOL 17200), Dec. 2022; 5 ♂♂ (9.1–12.7 mm) (NCHUZOOL 17201), Dec. 2022. C. purpureus Stimpson, 1858: Japan (purchased): 1 ♂, 22.1 mm (NCHUZOOL 17215), 1 ♂ (21.6) mm (NCHUZOOL 17216), 1 ♀, 18.6 mm (NCHUZOOL 17217). C. scaevola (Forskål, 1775): Djibouti: 3 ♂♂ (8.3–10.2 mm), 2 ♀♀ (7.9, 9.0 mm) (UF 33344), Moucha Islands, Maskali Bank, Sunken Buoy dive site, 27 Sep. 2012; Oman: 2 ♂♂ (17.5, 17.9 mm), 5 ♀♀ (14.8–18.4 mm) (UF 7609), Bar al Hikman peninsula, E side shore of peninsula, 19 Jan. 2005; Iran: 1 ♂ (16.8 mm), 1 ♀ (11.9 mm) (ZRC 2017.0642), Djod Village, Gulf of Oman, coll. S. Ebrahimnezhad, Oct. 2014. RESULTS TAXONOMY Family Coenobitidae Dana, 1851 Genus Coenobita Latreille, 1829 Coenobita moluccensis n. sp. (Figs. 1‒4, 13A, B) urn:lsid:zoobank.org:act:6591BEEE-2971-41AF-909F192976A23B26 Material examined: Holotype: ♂ (12.0 mm) (MZB Cru 5722), Aru Island, Maluku, coll. local Table 1. Haplotypes of 16S rDNA and cytochrome c oxidase subunit I (COI) genes of Coenobita species used in this study. *, holotype; **, paratype. See MATERIALS AND METHODS for abbreviations of museums and universities Species Locality Catalog no. (or tissue sample [TS] code) Sample size Haplotype of 16S Access. no. of 16S Haplotype of COI Access. no. of COI C. brevimanus Taiwan: Dongsha I. NCHUZOOL 17212 1 Cbr PQ427321 Cbr_C1 OR413746 Philippines: Olango I., Cebu UF 3799 1 Cbr PQ427322 Cbr_C2 OR413747 Thailand: Similan ZRC SM-03 1 Cbr PQ427323 Cbr_C3 OR413748 C. cavipes Taiwan: Lyudao, Taitung NCHUZOOL 13625 1 Cca1 PQ427324 Cca_C1 AB998653 Taiwan: Lyudao, Taitung NCHUZOOL 13626 1 Cca1 PQ427325 Cca_C2 AB998654 Taiwan: Houwan, Kenting NCHUZOOL 13627 1 Cca2 PQ427326 Cca_C3 AB998655 Taiwan: Yuanjhonggang, Kaohsiung NCHUZOOL 13628 1 Cca1 PQ427327 Cca_C4 AB998656 Philippines: Kawasan Falls, Cebu UF 11345 1 Cca3 PQ427328 Cca_C5 AB998657 C. lila Singapore: St John’s I. NCHUZOOL 13635 1 Cli1 PQ427329 Cli_C1 AB998648 Singapore: Pulau Jong NCHUZOOL 13636 1 Cli2 PQ427330 Cli_C2 AB998649 C. longitarsis Papua New Guinea UF 11434 1 Clo1 PQ427331 Clo_C1 OR413749 Papua New Guinea UF 11438 1 Clo2 PQ427332 Clo_C2 OR413750 Papua New Guinea UF 11435 1 Clo3 PQ427333 Clo_C3 OR413751 Papua New Guinea UF 11436 1 Clo3 PQ427334 Clo_C4 OR413752 page 3 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan Species Locality Catalog no. (or tissue sample [TS] code) Sample size Haplotype of 16S Access. no. of 16S Haplotype of COI Access. no. of COI C. perlatus Guam NCHUZOOL 17213 1 Cpe PQ427335 Cpe_C1 OR413753 Christmas Island ZRC CI-D07-2011 1 Cpe PQ427336 Cpe_C2 OR413754 Taiwan: Taiping I. NCHUZOOL 17214 1 Cpe PQ427337 Cpe_C3 OR413755 C. pseudorugosus Philippines: Panglao, Bohol UF 13164 1 Cps1 PQ427338 Cps_C1 OR413757 Philippines: Camiguin NCHUZOOL 17197 1 Cps1 PQ427339 Cps_C2 OR413760 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 17201 1 Cps2 PQ427340 Cps_C3 OR413768 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 17200 1 Cps1 PQ427341 Cps_C4 OR413770 C. purpureus purchased NCHUZOOL 17215 1 Cpu1 PQ427342 Cpu_C1 OR413775 purchased NCHUZOOL 17216 1 Cpu2 PQ427343 Cpu_C2 OR413776 purchased NCHUZOOL 17217 1 Cpu3 PQ427344 Cpu_C3 OR413777 C. rugosus Taiwan: Taiping I. NCHUZOOL 17204 1 Cru1 PQ427345 Cru_C1 OR413778 Philippines: Camiguin NCHUZOOL 17218 1 Cru2 PQ427346 Cru_C2 OR413779 Christmas I. ZRC 1 Cru3 PQ427347 Cru_C3 OR413780 C. spinosus Niue UF 2449 1 Csp1 PQ427348 Csp_C1 OR413781 Vanuatu UF 7501 1 Csp2 PQ427349 Csp_C2 OR413782 Wallis and Futuna NCHUZOOL 17216 1 Csp2 PQ427350 Csp_C3 OR413783 C. variabilis Australia: West Governor I., WA QM W21232 1 Cva1 PQ427351 Cva_C1 PQ426898 Australia: West Governor I., WA QM W21232 1 Cva2 PQ427352 Cva_C2 PQ426899 Australia: West Governor I., WA QM W21232 1 Cva2 PQ427353 Cva_C3 PQ426900 Indonesia: West Papua NCHUZOOL 15257 2 Cva3 PQ427354, PQ427355 Cva_C4 PQ426901, PQ426902 Indonesia: West Papua NCHUZOOL 15258 2 Cva3 PQ427356, PQ427357 Cva_C5 PQ426903, PQ426904 C. violascens Taiwan: Yuanjhonggang, Kaohsiung NCHUZOOL 13629 1 Cvi PQ427358 Cvi_C1 AB998658 Taiwan: Yuanjhonggang, Kaohsiung NCHUZOOL 13630 1 Cvi PQ427359 Cvi_C2 AB998659 Taiwan: Dongsha I., Kaohsiung NCHUZOOL 13631 1 Cvi PQ427360 Cvi_C3 AB998660 Taiwan: Dongsha I., Kaohsiung NCHUZOOL 13632 1 Cvi PQ427361 Cvi_C4 AB998661 Philippines: Kawasan Falls, Cebu UF 11347 1 Cvi PQ427362 Cvi_C5 AB998664 C. moluccensis Indonesia: Aru, Maluku (TS COx94) 1 Cmo1 PQ427363 Cmo_C1 PQ426905 Indonesia: Aru, Maluku (TS COx100) 1 Cmo2 PQ427364 Cmo_C2 PQ426906 Indonesia: Aru, Maluku NCHUZOOL 15259** 1 Cmo2 PQ427365 Cmo_C2 PQ426907 Indonesia: Aru, Maluku (TS COx101) 1 Cmo3 PQ427366 Cmo_C4 PQ426908 Indonesia: Aru, Maluku (TS COx102) 1 Cmo4 PQ427367 Cmo_C5 PQ426909 Indonesia: Aru, Maluku (TS COx103) 1Cmo5 PQ427368 Cmo_C6 PQ426910 Indonesia: Aru, Maluku (TS COx104) 1Cmo6 PQ427369 Cmo_C7 PQ426911 Indonesia: Aru, Maluku (TS COx105) 1 Cmo7 PQ427370 Cmo_C8 PQ426912 Indonesia: Aru, Maluku MZB Cru 5722* 1 Cmo7 PQ427371 Cmo_C8 PQ426913 Indonesia: Aru, Maluku (TS COx106) 1 Cmo4 PQ427372 Cmo_C9 PQ426914 Indonesia: Aru, Maluku MZB Cru 5723** 1 Cmo4 PQ427373 Cmo_C9 PQ426915 Indonesia: Aru, Maluku MZB Cru 5723** 1 Cmo4 PQ427374 Cmo_C9 PQ426916 Indonesia: Aru, Maluku ZRC 2023.0252** 1 Cmo4 PQ427375 Cmo_C9 PQ426917 Indonesia: Aru, Maluku MZB Cru 5723** 1 Cmo2 PQ427376 Cmo_C10 PQ426918 Indonesia: Aru, Maluku ZRC 2023.0252** 1Cmo9 PQ427377 Cmo_C11 PQ426919 Indonesia: Aru, Maluku NCHUZOOL 15259** 1Cmo8 PQ427378 Cmo_C3 PQ426920 C. patsyae Indonesia: Central Sulawesi (TS COx87) 1 Cpa1 PQ427379 Cpa_C1 PQ426921 Indonesia: Central Sulawesi (TS COx88) 1 Cpa2 PQ427380 Cpa_C2 PQ426922 Indonesia: Central Sulawesi (TS COx89) 1 Cpa1 PQ427381 Cpa_C3 PQ426923 Indonesia: Central Sulawesi (TS COx90) 1 Cpa3 PQ427382 Cpa_C4 PQ426924 Indonesia: Central Sulawesi (TS COx91) 1 Cpa4 PQ427383 Cpa_C5 PQ426925 Indonesia: Central Sulawesi (TS COx92) 1Cpa5 PQ427384 Cpa_C6 PQ426926 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5725* 1Cpa6 PQ427385 Cpa_C7 PQ426927 Indonesia: Central Sulawesi NCHUZOOL 15271 1Cpa6 PQ427386 Cpa_C7 PQ426928 Table 1. (Continued) page 4 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan Species Locality Catalog no. (or tissue sample [TS] code) Sample size Haplotype of 16S Access. no. of 16S Haplotype of COI Access. no. of COI Indonesia: Southeast Sulawesi NCHUZOOL 15272 2Cpa6 PQ427387, PQ427388 Cpa_C7 PQ426929, PQ426930 Indonesia: Central Sulawesi NCHUZOOL 15275 1Cpa6 PQ427389 Cpa_C7 PQ426931 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5726** 1 Cpa2 PQ427390 Cpa_C8 PQ426932 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2023.0254** 1 Cpa2 PQ427391 Cpa_C9 PQ426933 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2023.0254** 1 Cpa2 PQ427392 Cpa_C10 PQ426934 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2023.0254** 1Cpa8 PQ427393 Cpa_C11 PQ426935 Indonesia: Central Sulawesi NCHUZOOL 15271 1Cpa9 PQ427394 Cpa_C12 PQ426936 Indonesia: Central Sulawesi NCHUZOOL 15271 1 Cpa10 PQ427395 Cpa_C13 PQ426937 Indonesia: Central Sulawesi NCHUZOOL 15271 1 Cpa7 PQ427396 Cpa_C14 PQ426938 Indonesia: Central Sulawesi NCHUZOOL 15271 1 Cpa11 PQ427397 Cpa_C15 PQ426939 Indonesia: Southeast Sulawesi NCHUZOOL 15272 1 Cpa12 PQ427398 Cpa_C16 PQ426940 Indonesia: Southeast Sulawesi NCHUZOOL 15272 1 Cpa13 PQ427399 Cpa_C17 PQ426941 Indonesia: Southeast Sulawesi NCHUZOOL 15272 1 Cpa13 PQ427400 Cpa_C15 PQ426942 Indonesia: Southeast Sulawesi NCHUZOOL 15272 1 Cpa14 PQ427401 Cpa_C18 PQ426943 Indonesia: Southeast Sulawesi NCHUZOOL 15272 1 Cpa7 PQ427402 Cpa_C19 PQ426944 Indonesia: Central Sulawesi NCHUZOOL 15275 1 Cpa7 PQ427403 Cpa_C19 PQ426945 Indonesia: Central Sulawesi NCHUZOOL 15273 1Cpa15 PQ427404 Cpa_C20 PQ426946 Indonesia: Southeast Sulawesi NCHUZOOL 15274 1Cpa6 PQ427405 — Indonesia: Central Sulawesi NCHUZOOL 15275 1Cpa6 PQ427406 Cpa_C21 PQ426947 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15278 1 Cpa1 PQ427407 Cpa_C22 PQ426948 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15278 1 Cpa2 PQ427408 Cpa_C23 PQ426949 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15278 1 Cpa2 PQ427409 Cpa_C24 PQ426950 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15278 1 Cpa1 PQ427410 Cpa_C25 PQ426951 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15280 1 Cpa1 PQ427411 Cpa_C26 PQ426952 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15280 1 Cpa1 PQ427412 Cpa_C27 PQ426953 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15280; MZB Cru 5726** 2 Cpa7 PQ427413, PQ427414 Cpa_C28 PQ426954, PQ426955 C. celebensis Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5727* 1 Cce1 PQ427415 Cce_C1 PQ426956 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2024.0263** 1 Cce1 PQ427416 Cce_C1 PQ426957 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5728** 1 Cce1 PQ427417 Cce_C2 PQ426958 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5728** 1 Cce1 PQ427418 Cce_C3 PQ426959 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2024.0263** 1 Cce1 PQ427419 Cce_C4 PQ426960 Indonesia: Central Sulawesi NCHUZOOL 15276** 1 Cce2 PQ427420 Cce_C5 PQ426961 Indonesia: Central Sulawesi NCHUZOOL 15277** 1 Cce1 PQ427421 Cce_C6 PQ426962 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15279** 1 Cce1 PQ427422 Cce_C7 PQ426963 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15281** 1 Cce1 PQ427423 Cce_C8 PQ426964 C. granularis Indonesia: Central Sulawesi (TS COx95) 1 Cgr1 PQ427424 Cgr_C1 PQ426965 Indonesia: Central Sulawesi (TS COx96) 1 Cgr1 PQ427425 Cgr_C2 PQ426966 Indonesia: Central Sulawesi (TS COx97) 1 Cgr1 PQ427426 Cgr_C3 PQ426967 Indonesia: Central Sulawesi (TS COx98) 1 Cgr1 PQ427427 Cgr_C4 PQ426968 Indonesia: Central Sulawesi (TS COx99) 1 Cgr2 PQ427428 Cgr_C5 PQ426969 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5729* 1 Cgr3 PQ427429 Cgr_C6 PQ426970 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5730** 1Cgr5 PQ427430 Cgr_C7 PQ426971 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5791 1Cgr5 PQ427431 Cgr_C6 PQ426972 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5791 1 Cgr4 PQ427432 Cgr_C8 PQ426973 Indonesia: Central Sulawesi: Gulf of Tomini MZB Cru 5791 1 Cgr1 PQ427433 Cgr_C9 PQ426974 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2023.0253 1Cgr5 PQ427434 Cgr_C10 PQ426975 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2023.0253 1Cgr5 PQ427435 Cgr_C11 PQ426976 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2024.0264** 1 Cgr1 PQ427436 Cgr_C12 PQ426977 Indonesia: Central Sulawesi: Gulf of Tomini ZRC 2024.0264** 1 Cgr1 PQ427437 Cgr_C13 PQ426978 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15254 1 Cgr1 PQ427438 Cgr_C14 PQ426979 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15254 1 Cgr1 PQ427439 Cgr_C15 PQ426980 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15254 1 Cgr1 PQ427440 Cgr_C16 PQ426981 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15255** 1 Cgr1 PQ427441 Cgr_C17 PQ426982 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15256 1 Cgr1 PQ427442 Cgr_C18 PQ426983 Indonesia: Central Sulawesi: Gulf of Tomini NCHUZOOL 15256 1Cgr1 PQ427443 Cgr_C19 PQ426984 Table 1. (Continued) page 5 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan people, Dec. 2022. Paratypes: 2 ♂♂ (9.9, 14.6 mm), 1 ovig. ♀ (6.7 mm) (MZB Cru 5723), 3 ♂♂ (9.3, 11.3, 16.0 mm) (ZRC 2023.0252), 3 ♂♂ (10.5, 11.7, 14.4 mm) (NCHUZOOL 15259), same data as holotype. Diagnosis: Shield (Figs. 1A, 4A‒D) transversely convex, about 1.6 times as long as broad; dorsal surface with scattered small, flattened tubercles. Ocular peduncles reaching half length of fifth article of antennal peduncles; antennal peduncles reaching half length of penultimate article of antennular peduncles; antennular basal article 0.7 times as long as penultimate article (Fig. 1B); second article of antennal peduncle (Figs. 1C, 13A, B) stout, short. Chelipeds (Fig. 2A, B) unequal, dissimilar; left palm without stridulatory apparatus on upper outer surface; upper margin of palm of left and right cheliped with brush of long, coarse setae on proximal half; outer surface of palm of left cheliped with rows of tubercles, large and closelyset on upper half, smaller and fewer on lower half, some accompanied by tuft of very short setae; lower proximal part angled, forming broadly triangular lobelike projection, continued to slightly upright lower margin of fixed finger. P2 and P3 dissimilar, P2 slightly shorter, more slender than P3; dactylus and propodus of left P2 (Fig. 3A, B) with small, sometimes corneoustipped tubercles; ventral surface of dactylus with longitudinal ridge consisting of row of tiny corneous teeth; dactylus and propodus of right P2 covered with rows of corneous-tipped tubercles on lateral surface, larger tubercles on lower margin and ventral surface. Fig. 1. Coenobita moluccensis n. sp. Holotype ♂ (SL 12.0 mm, MZB Cru 5722). A, shield and cephalic appendages; B, right antennular peduncles, lateral view; C, right antennal peduncles, lateral view; D, sternite and coxae of male P5; E, telson. Setae partially omitted. Scale bars: A = 2 mm; B‒E = 1 mm. page 6 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan Left P3 (Fig. 3C–E) with dactylus slightly convex laterally, punctate, with scattered short tufts of setae; ventral surface with longitudinal ridge consisting of row of widely-spaced tiny corneous teeth; lateral surface of propodus broad, slightly flattened distally, punctate or with flattened small tubercles; dorsal surface broader distally; right P3 (Fig. 3F, G) with dactylus and propodus with tubercles on lateral surface, ventral surface of dactylus with scattered corneoustipped spines. P4 semi-chelate (Fig. 2C). In male, coxae of P5 (Figs. 1D, 2D) produced, thick, each forming moderately short, calcified sexual tube; moderately large, subquadrangular sternal protuberance between both coxae. Telson (Fig. 1E) with posterior lobes bearing very narrow median cleft. Description: Shield (Figs. 1A, 4A‒D) transversely convex, about 1.6 times as long as broad; anterior margin between rostrum and lateral projections slightly concave; lateral projection produced, terminating in pointed spine; posterior margin slightly rounded. Rostrum broadly rounded, obsolete. Lateral surface punctate; dorsal surface with scattered small, flattened tubercles. Ocular peduncles compressed laterally, mesial surface shallowly concave distally, reaching half length of fifth article of antennal peduncles; cornea small, occupying only one-third of distal part of ocular peduncles laterally; dorsal surface with scattered small, flattened tubercles and sparse short setae. Ocular acicles triangular, terminating acutely. Antennular peduncles (Fig. 1B) long. Basal article 0.7 times as long as penultimate article. Ultimate article longer than penultimate article. Upper rami of flagella stick-like, terminating in rounded tip, with very short setae on lateral and mesial margins, lower rami with 7 segments, long seta distally. Antennal peduncles (Figs. 1C, 13A, B) exceeding ocular peduncles by half length of fifth article, reaching Fig. 2. Coenobita moluccensis n. sp. Holotype ♂ (SL 12.0 mm, MZB Cru 5722). A, left cheliped, outer view; B, right cheliped, outer view; C, left P4, lateral view; D, sternite and coxae of male P5. page 7 of 38 Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan half length of penultimate article of antennular peduncles. First article 1.3 times as long as broad. Second article stout, short, not reaching proximal area of fourth article, covered with small tubercles and sparse very short setae. Third to fifth articles unarmed, with scattered, very short setae. Antennal acicle fused to second article of peduncle. Flagella long, overreaching tip of right cheliped. Chelipeds (Figs. 2A, B, 4A‒D) unequal, dissimilar; left distinctly larger than right. Dactylus of left cheliped (Fig. 2A) longer than palm; outer surface with numerous large tubercles, with tufts of short setae, upper surface with large, flattened tubercles, inner surface with rows of corneous-tipped tubercles and tufts of very short, stiff setae; cutting edge with 1 large tooth proximally and distally, smaller tooth medially, terminating in small corneous claw. Fixed finger with sparse, tubercles on outer surface, some tubercles with 2 or 3 short setae; lower margin with row of tubercles, some with tufts of short setae; cutting edge with 2 large teeth and row of smaller teeth, terminating in small corneous claw; inner surface covered with large tubercles, each with short setae. Palm without stridulatory apparatus on upper outer surface, upper margin with row of large, tubercles, brush of long, coarse setae on proximal half; outer surface with rows of tubercles, large and closely-spaced on upper half, smaller and fewer on lower half, some accompanied by tuft of very short setae; lower margin with small, flattened tubercles bearing tufts of very short setae; lower proximal part angled, forming broadly triangular lobe-like projection, continued to slightly upright lower margin of fixed finger; inner surface with small and large tubercles, some with short and long setae, brush of setae proximally near upper margin. Carpus with sparse corneous-tipped tubercles on upper half of outer surfaces, each tubercle with tufts of setae, lower half smooth or with very low tubercles; inner surface with flattened tubercles, each tubercle accompanied with moderately long setae, long, setae on distal margin. Merus with transverse rows of flattened tubercles on outer surface; lower outer and inner margins each with tubercles and tufts of short setae, inner surface smooth. Right cheliped (Fig. 2B) with dactylus longer than palm, outer surface covered with large, tubercles, some tubercles with corneous-tipped, inner surface with row of corneous-tipped tubercles and moderately long setae; cutting edge with row of moderately large teeth, terminating in small corneous claw. Fixed finger with cutting edge bearing several small teeth, large teeth adjacent terminal corneous claw and another in middle. Palm with thick brush of long coarse setae on upper margin; outer surface with rows of large tubercles, some with corneous-tipped, each with 2 or 3 long setae. Inner surface with scattered flattened tubercles, each with tufts of setae; proximal area of palm with long, coarse setae near upper margin. Carpus and merus similarly armed as left cheliped, but with long setae on lower margin of merus. P2 and P3 dissimilar, P2 slightly shorter, more slender than P3. Dactylus of left P2 with lateral and dorsal surfaces flattened, covered with rows of corneous-tipped, tubercles, each with short stiff setae (Fig. 3A); mesial surface with rows of numerous corneous-tipped tubercles each bearing tufts of short setae; ventral surface (Fig. 3B) with longitudinal ridge consisting of row of tiny corneous teeth, ventrolateral margin with row of corneous-tipped tubercles and tufts of short setae. Propodus lateral surface covered with dense, small, sometimes corneous-tipped tubercles, each with 1 or 2 setae; dorsal surface broad, with row of small corneous-tipped tubercles; dorsomesial margin with row of large, corneous-tipped tubercles; mesial surface concave, smooth except for few small tubercles and setae, delimited ventrally by longitudinal row of small tubercles. Carpus with rows of large tubercles on dorsolateral margin, each bearing 2 or 3 short setae; lateral surface with sparse, small, corneous-tipped tubercles, each with tufts of setae; ventrolateral margin with larger corneous-tipped tubercles each with tufts of long setae; mesial surface concave with scattered tufts of short setae; ventromesial surface with sparse tufts of short setae. Merus compressed laterally and mesially, lateral and mesial surfaces smooth, except for several tubercles distolaterally. Right P2 with dactylus covered with rows of corneous-tipped tubercles on lateral surface, larger tubercles on lower margin and ventral surface, dorsal margin not delimited, ventral surface with sparse corneous spines. Propodus, carpus and merus as in left P2. Left P3 (Fig. 3C) with dactylus 1.3–1.5 times as long as propodus, lateral surface slightly convex distally, flattened or slightly concave proximally, punctate but with scattered short tufts of setae, delimited dorsally by row of small tubercles, sometimes corneous-tipped, bearing short setae, dorsal surface with longitudinal rows of corneous-tipped tubercles each with tufts of setae; tubercles on dorsal margin and dorsal surface decreasing in size proximally; mesial surface with rows of tubercles each with tufts of setae; ventrolateral margin with row tubercles, with corneoustipped distally, each tubercles with long setae; ventral surface (Fig. 3D) slightly concave, median longitudinal ridge consisting of row of, widely-spaced, tiny corneous teeth. Propodus lateral surface broad, slightly flattened distally, slightly convex proximally, punctate or with flattened small tubercles; dorsolateral margin delimited page 8 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan distally by row of small tubercles; dorsal surface (Fig. 3E) flattened, broadened distally, slightly narrowing proximally, with rows of small, flattened tubercles, each with very short setae, dorsomesial margin delimited by row of low tubercles, each with short setae; mesial surface slightly concave, with sparse tubercles, some with corneous-tipped, each with tufts of setae; ventral surface slightly flattened distally, concave proximally, with irregular rows of corneous-tipped tubercles, each with tufts of long setae; ventrolateral margin with row of small tubercles and tufts of sparse setae. Carpus with rows of tubercles, decreasing in size proximally, each with 2 or 3 short setae on dorsolateral margin, lateral surface with numerous corneous-tipped tubercles each with tufts of setae; ventrolateral margin with moderately large spines each with tufts of setae; mesial surface compressed, almost smooth except for few tufts of short setae. Merus compressed laterally and mesially, transverse row of flattened tubercles on lateral surface; mesial surface almost smooth, ventral margin with moderately large spines and tufts of long setae. Right P3 more slender than left, with longer and denser setae on ventral margin. Dactylus and propodus (Fig. 3F, G) covered with tubercles, corneous-tipped tubercles mainly on dorsoand ventrolateral surfaces; dorsal margin not delimited; mesial surface of dactylus with rows of corneous-tipped tubercles and tufts of setae, ventral surface with sparse small corneous spines; mesial surface of propodus with sparse small tubercles and short setae. Carpus and merus as in left P3. P4 semi-chelate (Fig. 2C). Dactylus with row of small corneous teeth ventrally, long, coarse setae dorsally; propodal rasp well developed, occupying large, semicircular area, consisting of numerous corneous Fig. 3. Coenobita moluccensis n. sp. Holotype ♂ (SL 12.0 mm, MZB Cru 5722). A, dactylus and propodus of left P2, lateral view; B, dactylus of left P2, ventral view; C, dactylus and propodus of left P3, lateral view; D, dactylus of left P3, ventral view; E, propodus of left P3, dorsal view; F dactylus and propodus of right P3, lateral view; G, dactylus of right P3, ventral view. page 9 of 38 Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan triangular, lobe-like projection, continued to slightly upright lower margin of fixed finger; outer surface of fixed finger with large tubercles. Right cheliped covered entirely with corneous-tipped tubercles each with short setae. P2 and P3 dissimilar, left stouter than right; P2 shorter, more slender than P3; dactylus and propodus of P2 (Fig. 11A, B) with corneous-tipped tubercles on lateral surfaces, dorsal surface broad, ventral surface of left P2 with median longitudinal ridge consisting of row of tiny corneous teeth; ventral surface of right P2 with irregular rows of corneous teeth. Left P3 (Fig. 11C) with lateral surface of dactylus punctate or with sparse tubercles; delimited dorsally by row of small tubercles; ventral surface (Fig. 11D) slightly concave, with median longitudinal ridge consisting of row of tiny corneous teeth; propodus convex on lateral surface, covered with transverse rows of small tubercles; dorsolateral margin delimited by row of small tubercles; dorsal surface (Fig. 11E) flattened, slightly broadened distally, covered with rows of flattened tubercles, each with very short setae, dorsomesial margin delimited by row of low tubercles, each with short setae. Right P3 (Fig. 11F, G) with dactylus covered with large tubercles, lateral surface of propodus with dense, transverse rows of large tubercles, ventral surface of dactylus with irregular rows of corneous-tipped tubercles. P4 semi-chelate (Fig. 10C). In male, coxae of P5 thick, each forming moderately short, subtriangular sexual tube; large, strongly produced, ovate sternal protuberance between both coxae (Figs. 9D, 10D). Telson (Fig. 9E) with posterior lobes asymmetrical, left slightly longer; lobes separated by narrow median cleft, margins with row of setae. Fig. 10. Coenobita celebensis n. sp. Holotype ♂ (SL 18.3 mm, MZB Cru 5727). A, left cheliped, outer view; B, right cheliped, outer view; C, left P4, lateral view; D, sternite and coxae of male P5. page 16 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan Color in life (Fig. 12): Usually dark brown in adults. Surface of shield with patch of black, dark brown or light brown anteriorly, transverse dark brown line distally; sometimes broad longitudinal dark brown stripe medially, orangish yellow line posteriorly. Ocular peduncles orangish brown or light brown mesially; cornea black or dark brown. Antennal peduncles greenish brown, flagella orange, antennular peduncles orange. Chelipeds with merus bright yellow or orange. P2 and P3 with merus partially yellow or orange; each segment having black or dark brown patches. Size: Largest male SL 18.3 mm. Etymology: The specific name celebensis is derived from Celebes (= Sulawesi), where all specimens were collected and are distributed to date. Ecological notes: In Central Sulawesi, the habitat consists of sandy beaches bordered by rice fields and some creeks. This species is occasionally sympatric with C. violascens. Distribution: Indonesia (Central Sulawesi). Remarks: Coenobita moluccensis n. sp., C. patsyae n. sp., and C. celebensis n. sp. are very similar, sharing the following characters: palms of chelipeds covered with large and small tubercles; upper margin with brush of setae; no stridulatory ridge on the left cheliped palm, moderately short, triangular, stout male sexual tube on the coxae of P5; and yellow tinges on parts of the chelipeds and pereopods. Fig. 11. Coenobita celebensis n. sp. Holotype ♂ (SL 18.3 mm, MZB Cru 5727). A, dactylus and propodus of left P2, lateral view; B, dactylus of left P2, ventral view; C, dactylus and propodus of left P3, lateral view; D, dactylus of left P3, ventral view; E, propodus of left P3, dorsal view; F dactylus and propodus of right P3, lateral view; G, dactylus of right P3, ventral view. page 17 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan Fig. 12. Color in life of Coenobita celebensis n. sp. C, paratype ♂ (SL 15.5 mm, NCHUZOOL 15277, Central Sulawesi, Indonesia). A, B, D–F, specimens from Sulawesi, Indonesia; not kept. page 18 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan Fig. 13. Outer surface of the second article of the right antennal peduncles in Coenobita moluccensis n. sp. (A, B), C. patsyae n. sp. (C, D), C. celebensis n. sp. (E, F), and C. lila Rahayu, Shih & Ng, 2016 (G, H.). A, holotype ♂ (SL 12.0 mm; MZB Cru 5722); B, paratype ♂ (SL 14.4 mm; NCHUZOOL 15259); C, holotype ♂ (SL 15.0 mm; MZB Cru 5725); D, ♂ (SL 13.8 mm; NCHUZOOL 15273); E, holotype ♂ (SL 18.3 mm, MZB Cru 5727); F, paratype ♂ (SL 15.5 mm, NCHUZOOL 15277); G, paratype ♂ (SL 10.7 mm, NCHUZOOL 13624); H, ♂ (SL 11.3 mm NCHUZOOL 15284). Scale bars = 1 mm. page 19 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan These three new species can be distinguished by the following characters (Table 2): 1) morphology of left P3 propodus: in C. moluccensis, dorsolateral margin delimited by row of small tubercles, with basal lateral surface slightly convex; surfaces punctate in most specimens (Fig. 3C); in C. patsyae, dorsolateral margin not delimited or weakly delimited by row of small tubercles, with basal lateral surface slightly convex; surfaces covered with small tubercles in most specimens (Fig. 7C); and in C. celebensis, dorsolateral margin delimited by row of small tubercles, with basal lateral surface convex; surfaces covered with large tubercles in most specimens (Fig. 11C); 2) shape of second article of antennal peduncles: stout (ratio of length from tip to inner base along middle/width from dorsal margin to inner base about 1.4) and short (not reaching proximal area of fourth article) in C. moluccensis (Figs. 1C, 13A, B); slender (ratio about 1.7) and long (reaching half length of fourth article) in C. patsyae (Figs. 5C, 13C, D); and broad or stout (ratio about 1–1.4) and short (barely reaching proximal area of fourth article) in C. celebensis (Figs. 9C, 13E, F); and 3) shape of sternal protuberance between male fifth coxae: large, subquadrangular in C. moluccensis (Figs. 1D, 2D); strongly produced, ovate in C. patsyae (Figs. 5D, 6D); strongly produced, long, ovate, in C. celebensis (Figs. 9D, 10D). Fize and Serène (1955) stated that color alone is unreliable for distinguishing species within the genus Coenobita. However, the varying degree of yellow or orange, especially on the cheliped merus, can still be useful for differentiating most individuals of the three species from other congeners, particularly in larger individuals. In C. moluccensis, there is only a yellow or orange tinge (Fig. 4); in C. patsyae, the color ranges from yellow or orange to a yellow or orange tinge, with some individuals showing no yellow or orange tint (Fig. 8); and in C. celebensis, the color is consistently yellow or orange (Fig. 12). The three new species are similar to C. lila and C. cavipes Stimpson, 1858 in which the left cheliped palm is covered by tubercles, without stridulatory ridge on the dorsal surface of the palm, with row of dense, stiff setae on dorsal surface of palm, and stout and subtriangular sexual tubes (e.g., ca. 1.5–1.6 times as long as broad measured on mesial margin of right tube), but show differences as follows: 1) left P3 propodus broader, 1.6 times as long as broad in C. moluccensis, 1.8 times in C. patsyae and C. celebensis (1.9 times in C. lila and 2.3 Table 2. Comparison of characters and fresh colorations of Coenobita moluccensis n. sp., C. patsyae n. sp., C. celebensis n. sp., C. lila Rahayu, Shih & Ng, 2016, and C. cavipes Stimpson, 1858 Characters C. moluccensis C. patsyae C. celebensis C. lila C. cavipes Antennal peduncles Second article stout, short, not reaching proximal area of fourth article (Figs. 1C, 13A, B). Second article slender, long, reaching half length of fourth article (Figs. 5C, 13C, D). Second article stout or broad, short, barely reaching proximal area of fourth article (Figs. 9C, 13E, F). Second article stout, short, not reaching proximal area of fourth article (Rahayu et al. 2016: fig. 1C; Fig. 13G, H). Second article slender, long, reaching half length of fourth article (Rahayu et al. 2016: fig. 7C). Propodus of left P3 Broader (1.6 times as long as broad); dorsolateral margin delimited by row of small tubercles, basal lateral surface slightly convex; surfaces punctate (Fig. 3C). Slightly broader (1.8 times as long as broad); dorsolateral margin not delimited or weakly delimited by row of small tubercles, basal lateral surface slightly convex; surfaces covered with small tubercles (Fig. 7C). Slightly broader (1.8 times as long as broad); dorsolateral margin delimited by row of small tubercles, basal lateral surface convex; surfaces covered with large tubercles (Fig. 11C). Sightly narrower (1.9 times as long as broad); dorsolateral margin delimited only on 4/5 by weak tubercles, basal lateral surface slightly convex; surfaces covered with corneous-tipped tubercles (Rahayu et al. 2016: figs. 2E, 5A–C, 6A–C). Narrower (2.3 times as long as broad); dorsolateral margin delimited by row of corneous-tipped tubercles, , basal lateral surface slightly convex; surfaces only with few scattered tubercles (Rahayu et al. 2016: figs. 5D–F, 6D–F, 8E). Coxae of P5 Sternal protuberance large, subquadrate (Figs. 1D, 2D). Sternal protuberance moderately large, ovate, produced (Figs. 5D, 6D). Sternal protuberance large, ovate, strongly produced (Figs. 9D, 10D). Sternal protuberance large, subpentagonal (Rahayu et al. 2016: figs. 1D, 2G). Sternal protuberance moderately large, cylindrical (Rahayu et al. 2016: figs. 7D, 8G). Color on cheliped merus Only a yellow or orange tinge (Fig. 4). Ranges from yellow or orange to a yellow or orange tinge, with some individuals showing no yellow or orange tint (Fig. 8). Consistently yellow or orange (Fig. 12). No yellow or orange tinge; some with very faint tinge (Rahayu et al. 2016: figs. 10, 11A–C). Consistently brown or greenish brown (Rahayu et al. 2016: fig. 11D–F). page 20 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan times in C. cavipes) (Table 2); 2) sternal protuberance subquadrangular in C. moluccensis; produced and ovate in C. patsyae; and strongly produced, ovate and long in C. celebensis (subpentagonal in C. lila and cylindrical in C. cavipes) (Table 2); and 3) dactyli of right P2 and P3 with scattered corneous spines on ventral surface in the three new species and C. lila, but a row of tiny corneous teeth medially in C. cavipes. Coenobita granularis n. sp. (Figs. 14‒17) urn:lsid:zoobank.org:act:A21F7F9F-761E-4072-AF49E1A49EE23A0B Material examined: Holotype: ♂ (10.4 mm) (MZB Cru 5729), Gulf of Tomini, Central Sulawesi, Indonesia, coll. local people, 2022. Paratypes: 1 ♂ (8.7 mm), 1 ♀ (10.9 mm) (MZB Cru 5730), 2 ♂♂ (10, 10.8 mm) (ZRC 2024.0263), 1 ♂ (10.7 mm), 1 ♀ (8.0 mm) (NCHUZOOL 15255), same data as holotype. Others: 4 ♂♂ (7.6, 8.7, 9.2, 11.5 mm), 1 ♀ (7.7 mm) (MZB Cru 5791), 4 ♂♂ (7.3, 8.3, 8.9, 11.1 mm), 1 ♀ (7.4 mm) (ZRC 2023.0253), 3 ♂♂ (8.7, 8.8, 9.0 mm) (NCHUZOOL 15254), 4 ♂♂ (8.2, 9.1, 9.6, 9.8 mm), 2 ♀♀ (8.5, 9.0 mm) (NCHUZOOL 15256), same data as holotype. Diagnosis: Shield (Figs. 14A, 17A‒C), transversely convex, about 1.5 times as long as broad; dorsal surface punctate, tubercles on dorsodistal surface. Ocular peduncles reaching third proximal of ultimate article of antennal peduncles, basal article of antennular peduncles 0.6 times as long as penultimate article (Fig. 14B); antennal peduncles (Fig. 14C) exceeding ocular peduncles by half length of fifth article, second article slender, smooth. Chelipeds (Fig. 15A, B) unequal and dissimilar, left larger than right, brush of setae on half proximal of upper margin of both palms; left cheliped palm with row of 5‒6 stridulatory apparatus (laminar ridge) on upper outer surface, outer surface with widely-spaced tubercles, sometimes corneous-tipped; lower margin of palm oblique, with corneous-tipped tubercles bearing tufts of short setae; lower proximal angled rounded or 3 cornered. P2 and P3 dissimilar (Fig. 16), P2 markedly more slender than P3. Lateral surfaces of dactylus and propodus of left P2 (Fig. 16A) with numerous flattened tubercles bearing tufts of short setae, ventral surface of dactylus (Fig. 16B) with longitudinal ridge consisting of row of tiny corneous teeth, closely-spaced proximally, widely-spaced distally. Right P2 slightly shorter, armament similar. Lateral surface of dactylus and propodus of left P3 (Fig. 16C–E) broad, punctate; ventral surface of dactylus (Fig. 16D) slightly concave, median longitudinal ridge consisting of row of tiny corneous teeth. Right P3 slightly more slender than left. Dactylus and propodus (Fig. 16F) covered with flattened tubercles; ventral surface of dactylus (Fig. 16G) with sparse, low corneous-tip tubercles. P4 semi-chelate (Fig. 15C). In male, coxae of P5 thick, each forming calcified sexual tube, right tube long, directed to left, left tube short, broader than right; gonopore positioned on moderately broad with truncate tip posterior projection or papillae, with short, sparse, setae; cylindrical, small, sternal protuberance between both coxae (Figs. 14D, F, 15D, E). Telson (Fig. 14E) posterior lobes slightly asymmetrical separated by narrow median cleft; margins rounded, with row of setae. Description: Shield (Figs. 14A, 17A‒C) transversely convex, about 1.5 times as long as broad; anterior margin between rostrum and lateral projections slightly concave; lateral projection produced, terminating in pointed spine; posterior margin slightly convex. Rostrum broadly rounded, obsolete. Lateral surface smooth, with numerous tufts of long setae; dorsal surface punctate, tubercles on dorsodistal surface. Ocular peduncles (Fig. 14A) concave laterally, mesial surface compressed, reaching third proximal of ultimate segment of antennal peduncles; cornea small, occupying slightly more than half of distal part of ocular peduncles laterally; dorsal surface punctate with short setae. Ocular acicles triangular, terminating acutely. Antennular peduncles (Fig. 14B) long. Basal segment 0.6 times as long as penultimate segment. Ultimate segment 1.1 times as long as penultimate segment. Upper rami of flagella stick-like, terminating in rounded tip, with very short setae on lateral and mesial margins. Antennal peduncles (Fig. 14C) exceeding ocular peduncles by half length of fifth segment, reaching third proximal of penultimate segment of antennular peduncles. First segment 1.4 times as long as broad. Second segment slender, smooth. Third, fourth and fifth segments unarmed, with scattered, very short setae. Antennal acicle short, fused to second segment of peduncle. Flagella long, far exceeding tip of left cheliped. Chelipeds (Fig. 15A, B) unequal and dissimilar, left larger than right. Dactylus of left cheliped (Fig. 15A) with numerous broad, flattened tubercles bearing short setae on outer surface, upper margin with row of corneous-tipped spines and short, stiff setae; inner surface with rows of corneous-tipped tubercles bearing short setae; cutting edge with 3 large teeth terminating in small corneous claw. Row of 5‒6 stridulatory apparatus (laminar ridge) on upper outer surface of palm. Outer surface of fixed finger with few tubercles, some with corneous-tipped; lower margin straight with row of large tubercles, some with corneous-tipped, each page 21 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan bearing tufts of short setae; inner surface with few of corneous tipped tubercles; cutting edge with 2 large and 2 small teeth, terminating in small corneous claw. Palm with upper margin bearing row of corneous spines and brush of long, coarse setae, outer surface with widelyspaced tubercles, sometimes corneous-tipped; lower margin oblique, with corneous-tipped tubercles bearing tufts of short setae; lower proximal angled rounded or 3 cornered; inner surface with punctate with short setae, longitudinal ridge consisted of large tubercle starting at base of cutting edge to articulation with carpus, larger tubercles adjacent to ridge; brush of setae proximally near upper margin. Carpus with tubercles with tufts of short setae arranged more or less in longitudinal rows on outer surface near upper margin, narrow, smooth area medially, less dense, flattened tubercles near lower margin; inner surface with few small, flattened tubercles bearing short tufts of setae. Merus with transverse rows of flattened tubercles bearing very short setae on outer surface; lower margin produced distally, lower inner margin almost smooth but with tufts of long setae; inner and lower surfaces smooth but with few transverse tufts of very short setae. Right cheliped (Fig. 15B) with upper margin of dactylus bearing row of corneous-tipped tubercles, outer surface with widely-spaced flattened tubercles and short setae; palm with row of corneous spines and thick brush of long coarse setae on upper margin; cutting edges of Fig. 14. Coenobita granularis n. sp. A‒E, holotype ♂ (SL 10.4 mm, MZB Cru 5729); F, paratype ♂ (SL 10.7 mm, NCHUZOOL 15255). A, shield and cephalic appendages; B, right antennular peduncles, lateral view; C, right antennal peduncles, lateral view; D, F, sternite and coxae of male P5; E, telson. Setae partially omitted. Scale bars: A = 2 mm; B‒E = 1 mm. page 22 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan dactylus and fixed finger each with large tooth adjacent to corneous claw, followed by row of small teeth. Outer surfaces of palm and fixed finger compressed, with rows of flattened, corneous-tipped tubercles, each bearing short setae, lower margin with row of long, stiff setae; inner surfaces of palm and dactylus with scattered corneous-tipped tubercles, each with tufts of setae; proximal area of palm with long, coarse setae near upper margin. Carpus and merus compressed laterally, similarly armed as left cheliped. P2 and P3 dissimilar (Fig. 16A, C, F), P2 markedly more slender than P3. Dactylus of left P2 with rows of corneous-tipped tubercles each with short setae on lateral surface; dorsal surface flattened, not delimited, with widely-spaced rows of corneoustipped tubercles; mesial surface with rows of widelyspaced flattened, corneous-tipped tubercles each bearing tufts of short setae; ventral surface (Fig. 16B) with longitudinal ridge consisting of row of tiny corneous teeth, closely-spaced proximally, widely-spaced distally, ventrolateral margin with row of corneous-tipped tubercles bearing tufts of setae. Propodus broad dorsal surface with rows of low tubercles, some with corneoustipped, dorsomesial margin with row of corneous-tipped tubercles, dorsolateral surface not delimited, lateral surface compressed, with numerous flattened tubercles bearing tufts of short setae; ventrolateral surface with row of corneous-tipped tubercles and rather long setae, mesial surface concave, transverse row of flattened tubercles each with short setae. Carpus with row of small tubercles each bearing short setae on dorsolateral margin, lateral surface with row of flattened tubercles each bearing tufts of setae; mesial surface compressed, with few tufts of short setae. Merus compressed laterally Fig. 15. Coenobita granularis n. sp. A‒D, holotype ♂ (SL 10.4 mm, MZB Cru 5729); E, paratype ♂ (SL 10.7 mm, NCHUZOOL 15255). A, left cheliped, outer view; B, right cheliped, outer view; C, left P4, lateral view; D, E, sternite and coxae of male P5. page 23 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan and mesially, with tufts of long setae on dorsal and ventral margins, transverse row of transverse tufts of setae on lateral surface; mesial surface almost smooth, ventral margin with row of moderately large tubercles. Right P2 slightly shorter than left. Armament similar. Left P3 with dactylus slightly shorter than or same length as propodus (Fig. 16C), lateral surface slightly convex distally, flattened proximally, punctate, delimited dorsally by row of tubercles bearing short setae, dorsal surface with longitudinal rows of corneous-tipped tubercles each with tufts of setae, decreasing in size proximally; mesial surface with rows of corneous-tipped tubercles each with tufts of setae; ventrolateral margin with row of widely-spaced corneous small spines with short setae; ventral surface (Fig. 16D) slightly concave, median longitudinal ridge consisting of row of tiny corneous teeth, not reaching tip. Propodus slightly flattened distally on lateral surface, becoming convex proximally, punctate; dorsolateral margin not delimited by row of tubercles; dorsal surface (Fig. 16E) flattened, distally broad, slightly narrower proximally, with rows of flattened tubercles; dorsomesial margin delimited by row of tubercles sometimes with corneous-tipped and short setae; mesial surface slightly concave, with transverse rows of tufts of setae; ventral surface slightly concave with widely-spaced tubercles and scattered tufts of setae; ventrolateral margin with row of tubercles and tufts of setae. Carpus with row of small tubercles each with short setae on dorsolateral margin, lateral surface with sparse low tubercles and tufts of setae; Fig. 16. Coenobita granularis n. sp. Holotype ♂ (SL 10.4 mm, MZB Cru 5729). A, dactylus and propodus of left P2, lateral view; B, dactylus of left P2, ventral view; C, dactylus and propodus of left P3, lateral view; D, dactylus of left P3, ventral view; E, propodus of left P3, dorsal view; F dactylus and propodus of right P3, lateral view; G, dactylus of right P3, ventral view. page 24 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan mesial surface compressed, smooth except for some tufts of setae. Merus compressed laterally and mesially, transverse row of flattened tubercles on lateral surface; mesial surface almost smooth, tufts of long setae on dorsal and ventral margin. Right P3 slightly more slender than left. Dactylus and propodus (Fig. 16F) covered by with flattened tubercles; mesial and ventral surfaces of dactylus (Fig. 16G) with low corneous-tip tubercles; mesial surface of propodus with transverse rows of corneous-tip small tubercles. Carpus and merus as in left P3. P4 semi-chelate (Fig. 15C), dactylus with row of small corneous teeth ventrally, long, coarse setae dorsally; propodal rasp well developed, occupying large, circular area, consisting of numerous corneous scales; propodus with distodorsal spine. P5 chelate. In male, coxae of P5 thick, each forming calcified sexual tube, right tube long, directed to left, left tube short, broader than right. Gonopore positioned on moderately broad with truncate tip posterior projection or papillae, with short, sparse, setae; cylindrical, small, sternal protuberance between both coxae (Figs. 14D, F, 15D, E). In females, coxae of P5 thick, subquadrate. Telson (Fig. 15E) with distinct lateral indentation, separating anterior and posterior lobes. Anterior lobe as long as posterior. Posterior lobes slightly asymmetrical separated by narrow median cleft; margins rounded, with row of setae. Variation: Morphological variation related to sex is observed in the degree of tuberculation and the density of setae on the chelipeds and ambulatory legs. In one female specimen (MZB Cru 5730), the tubercles on the ambulatory legs are larger and more pronounced, and the setae are denser. The left sexual tube of the holotype forms a sharp corner laterally near the truncated tip (Fig. 14D), while in the paratypes, the lateral margin is oblique near the truncated tip (Fig. 14F). It is possible that the holotype sexual tube is damaged or worn out, as it is also devoid of setae. Color in life: General light brown, orangish brown or dark brown (Fig. 17). Shield light brown or orangish brown with streak of green, darker brown dorsodistally. Ocular peduncles green or dark brown, corneas black. Antennal peduncles dark brown, with fifth segment dark orange. Antennular peduncles dark brown, with penultimate segment dark orange. Chelipeds with palms and dactyli light brown or orangish brown, cutting edges white; carpi and meri light brown or orangish brown. P2 and P3 with dactyli and propodi light brown; carpi and meri greenish brown or light brown tinge with dark brown. Size: Largest male SL 11.5 mm; largest female 10.9 mm. Etymology: The name is derived from the Latin word “granular,” meaning “small grain”, referring to the tuberculate surface of the chelipeds and pereopods in most specimens, especially in adults. Ecological notes: In Central Sulawesi, the habitat consists of sandy beaches with low shrubs, and the beach typically features a shallow intertidal zone. This species is sympatric with C. pseudorugosus (Table 1; Shih et al. 2023a) and C. rugosus, but individuals of C. granularis usually congregate separately from the other two species. Distribution: Indonesia (Gulf of Tomini, Central Sulawesi). Remarks: Among the species with row of a stridulatory ridge on the left cheliped palm in Coenobita, this new species most closely resembles C. perlatus, C. pseudorugosus, and C. purpureus Stimpson, 1858 in having distinctly unequal sexual tube with the right tube being more slender and much longer than the left. The morphological similarity between C. granularis n. sp. and C. pseudorugosus is also supported by molecular evidence (Fig. 23). In C. granularis and C. perlatus, the right sexual tube is 2 times as long as the left. However, the shape of the left sexual tube differs between the two species. In the new species, the left sexual tube is broad and stout, about 1.3 times as long as broad (measured along the mesial margin), and the sternal protuberance is small and ovate (Figs. 14D, F, 15D, E), while it is more slender, about 1.5 times as long as broad, sternal protuberance rounded, small in C. perlatus (Alcock 1905: pl. 14(2a); Nakasone, 1988: fig. 5F). In C. pseudorugosus and C. purpureus, the right sexual tube is shorter, about 1.6 and 1.2 times as long as the left respectively. The left sexual tube of C. pseudorugosus is 1.1 times as long as broad, the sternal protuberance is ovate and relatively small (Nakasone 1988: fig. 1H; Shih et al. 2023a: figs. 6B, 9A, C, E). In C. purpureus the left tube is 1.8 times as long as broad and the sternal protuberance is ovate and relatively large (Nakasone 1988: fig. 4F). Other differences observed include the left cheliped of C. granularis, which has a covering of flattened tubercles on the outer surface of the palm (Figs. 15A, 17D–F) [covered by large tubercles in C. perlatus (Nakasone 1988: fig. 5B) and C. pseudorugosus (Nakasone 1988: fig. 1C; Shih et al. 2023a: figs. 6E, 8A, C, E, G); with sparser tubercles on the outer upper surface and almost smooth on the lower outer surface in C. purpureus (Nakasone 1988: fig. 4B)]. The left P3 propodus is broad, with the dorsal margin not delimited by a row of tubercles; the lateral surface is slightly convex and punctate in C. granularis (Fig. 16C, E) [in C. perlatus, the dorsolateral margin of the P3 propodus is not delimited, and the lateral surface is convex and page 25 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan distances (and bp differences) are 0–3.3% (0–21 bp) within species, and 3.6–17.39% (23–100 bp) between species. DISCUSSION Integrative taxonomy and genetic divergence Species in the genus Coenobita have long been traded as ornamental crustaceans. They are attractive to hobbyists due to their beautiful coloration and the ease of keeping them alive in a terrarium (Bundhitwongrut 2018; Schäfer 2020). However, detailed taxonomic studies of this genus are limited. In the last 50 years, only two new species have been described, Coenobita pseudorugosus Nakasone, 1988 and C. lila Rahayu, Shih & Ng, 2016, with molecular support available for the latter. The identities of C. pseudorugosus and C. longitarsis have recently been confirmed by both morphological and molecular evidence (Shih et al. 2023a). Integrative taxonomy has become crucial in delineating species because it utilized multiple Fig. 22. Coenobita scaevola (Forskål, 1775), ♂ (SL 16.8 mm, ZRC 2017.0642). A, dactylus and propodus of left P2; B, merus of left cheliped, inner view; C, sternite and coxae of male P5; D, merus of right cheliped, inner view, showing the setal tuft (arrowed). page 32 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan Fig. 23. A maximum likelihood (ML) tree for Coenobita species from the Indo-West Pacific, based on the combined 16S rDNA and cytochrome c oxidase I genes. Probability values at the nodes represent support values for ML and Bayesian inference (BI). For haplotype names, see table 1. 0.05 Cpa1+Cpa_C22 Cpa1+Cpa_C3 Cpa4+Cpa_C5 Cmo2+Cmo_C2* (x2) Cmo1+Cmo_C1 Cpa5+Cpa_C6 Cgr1+Cgr_C2 Cbr+Cbr_C2 Cgr5+Cgr_C10 Cli2+Cli_C2 Cpa13+Cpa_C15 Cgr5+Cgr_C6 Cpe+Cpe_C1 Cmo9+Cmo_C11* Cpe+Cpe_C3 CfL2+CfL_C5* Cgr1+Cgr_C14 Cmo7+Cmo_C8** (x2) Cpa11+Cpa_C15 Clo3+Clo_C4 Cca3+Cca_C5 Cca1+Cca_C2 Cpa2+Cpa_C10* Cva2+Cva_C3 Cru1+Cru_C1 Cbr+Cbr_C3 Cvi+Cvi_C4 Cgr4+Cgr_C8 Cps1+Cps_C2 Cvi+Cvi_C2 Cpa2+Cpa_C2 Cpa3+Cpa_C4 Cmo2+Cmo_C10* CfL1+CfL_C4* Cps1+Cps_C1 Cru2+Cru_C2 Cpu2+Cpu_C2 Cpa6+Cpa_C21 Cpa15+Cpa_C20 Cgr1+Cgr_C19 CfL1+CfL_C6* Cgr1+Cgr_C13* Cgr1+Cgr_C16 Cvi+Cvi_C1 Cpa1+Cpa_C26 CfL1+CfL_C2* Cbr+Cbr_C1 Csp1+Csp_C1 CfL1+CfL_C8* Cgr1+Cgr_C18 Cmo4+Cmo_C9* (x4) Cpa7+Cpa_C19 (x2) Cgr2+Cgr_C5 Csp2+Csp_C3 Cpa2+Cpa_C24 Cva2+Cva_C2 Cca1+Cca_C4 Cgr1+Cgr_C12* Cmo8+Cmo_C3* Cgr1+Cgr_C15 Cpa1+Cpa_C25 Cva1+Cva_C1 Cpu1+Cpu_C1 Cpa10+Cpa_C13 Cpa13+Cpa_C17 Cpa9+Cpa_C12 Cpa2+Cpa_C9* CfL1+CfL_C1** (x2) Clo2+Clo_C2 Cca2+Cca_C3 Cpa2+Cpa_C8* Cca1+Cca_C1 Cpu3+Cpu_C3 Cpe+Cpe_C2 Cvi+Cvi_C5 Cva3+Cva_C4 (x2) Cgr5+Cgr_C7* Cpa1+Cpa_C27 Cmo4+Cmo_C5 Cgr3+Cgr_C6** Cpa12+Cpa_C16 Cpa6+Cpa_C7** (x5) Cli1+Cli_C1 Cmo3+Cmo_C4 CfL1+CfL_C7* Cpa7+Cpa_C14 Cpa2+Cpa_C23 Clo3+Clo_C3 Cvi+Cvi_C3 Cgr1+Cgr_C3 CfL1+CfL_C3* Csp2+Csp_C2 Cpa14+Cpa_C18 Cmo5+Cmo_C6 Cps1+Cps_C4 Cpa8+Cpa_C11* Cva3+Cva_C5 (x2) Clo1+Clo_C1 Cgr1+Cgr_C1 Cpa7+Cpa_C28 (x2) Cgr1+Cgr_C4 Cgr5+Cgr_C11 Cru3+Cru_C3 Cgr1+Cgr_C17* Cgr1+Cgr_C9 Cpa1+Cpa_C1 Cps2+Cps_C3 Cmo6+Cmo_C7 100/1 70/- 99/1 57/- 100/1 100/1 99/1 100/1 100/1 100/1 99/1 98/0.86 100/1 100/1 54/0.6 100/1 100/1 62/0.72 100/1 100/1 81/0.82 100/1 100/1 91/0.92 89/0.98 97/0.95 74/0.9 100/1 C. patsyae n. sp. C. celebensis n. sp. C. moluccensis n. sp. C. lila C. variabilis C. granularis n. sp. C. pseudorugosus C. cavipes C. spinosus C. violascens C. brevimanus C. rugosus C. purpureus C. longitarsis C. perlatus ML / BI support values page 33 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan characters, including DNA and various other data types, to delimit, discover, and identify meaningful, natural species and taxa at all levels (Dayrat 2005; Will et al. 2005; Padial et al. 2010; Goldstein and DeSalle 2011; Pante et al. 2015). Recent examples of integrative taxonomy in decapod crustaceans (e.g., Yuan et al. 2022; Shih et al. 2023c 2024). In our study, while the morphological differences among C. moluccensis, C. patsyae, and C. celebensis are considered minor (Table 2), molecular evidence plays an crucial role in distinguishing them, particularly regarding genetic distances (Table 3) and monophyly (Figs. 23, 24). The interspecific divergences of the DNA barcode gene COI among the species examined in this study are ≥ 3.6% (K2P distance) (Table 3). Within the species complex composed of Coenobita moluccensis, C. patsyae, and C. celebensis, the minimum interspecific divergences are smaller compared to values > 10% found in other coenobitid studies (e.g., 13.37% in Rahayu et al. 2016; 13.5% in Hamasaki et al. 2017; 10.18% in Shih et al. 2023a). However, these values are closer to the 4% range observed in the coastal hermit crabs Calcinus (Malay and Paulay 2010). Additionally, the three species are strongly supported in the phylogenetic tree (Fig. 23) and haplotype network (Fig. 24). As a result, they can be considered as pseudocryptic species (see Ng and Shih 2023; Shih et al. 2023a b c; Thurman et al. 2023) due to their similar morphology. Phylogeny and morphological and color traits Based on the phylogeny constructed from the Fig. 24. Haplotype network for the 16S+COI haplotypes observed from Coenobita patsyae n. sp., C. celebensis n. sp., and C. moluccensis n. sp. Unlabeled nodes represent inferred haplotypes not found in the sampled populations, and the hatch marks indicate the number of mutations. See table 1 for haplotype names. Cpa1+Cpa_C1 Cpa2+Cpa_C2 Cpa1+Cpa_C3 Cpa3+Cpa_C4 Cpa4+Cpa_C5 Cpa5+Cpa_C6 Cpa6+Cpa_C7 (x5) Cpa9+Cpa_C12 Cpa10+Cpa_C13 Cpa15+Cpa_C20 Cpa12+Cpa_C16 Cpa13+Cpa_C17 Cpa13+Cpa_C15 Cpa14+Cpa_C18 Cpa11+Cpa_C15 Cpa6+Cpa_C22 Cpa2+Cpa_C9 Cpa7+Cpa_C28 (x2) Cpa1+Cpa_C26 Cpa1+Cpa_C22 Cpa2+Cpa_C23Cpa2+Cpa_C24 Cpa2+Cpa_C10 Cpa1+Cpa_C25 COx154 Cpa1+Cpa_C27 Cpa8+Cpa_C11 Cpa2+Cpa_C8 Cpa7+Cpa_C19 (x2) CfL1+CfL_C7 CfL1+CfL_C1 (x2) CfL1+CfL_C8 CfL1+CfL_C2 CfL1+CfL_C3 CfL1+CfL_C4 CfL1+CfL_C6 CfL2+CfL_C5 Cmo1+Cmo_C1 Cmo2+Cmo_C2 (x2) Cmo2+Cmo_C10 Cmo3+Cmo_C4 Cmo4+Cmo_C5 Cmo5+Cmo_C6 Cmo6+Cmo_C7 Cmo7+Cmo_C8 (x2) Cmo4+Cmo_C9 (x4) Cmo8+Cmo_C3 Cmo9+Cmo_C11 C. celebensis C. patsyae C. moluccensis page 34 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan available sequences of the combined 16S and COI (Fig. 23), C. moluccensis, C. patsyae, and C. celebensis form a highly supported major clade that is sister to C. lila. Their close genetic relationship is also reflected in their similar morphology and coloration (Table 2), including the absence of a stridulatory ridge on the outer surface of the left chela (Figs. 2A, 4, 6A, 8, 10A, 12; Rahayu et al. 2016: figs. 2A, 4A–C, 10), the short sexual tubes in males (Figs. 1D, 2D, 5D, 6D, 9D, 10D; Rahayu et al. 2016: figs. 1D, 2G), and the yellow or orange tinges on the pereopods (see Remarks under C. celebensis). Coenobita patsyae and C. celebensis from Sulawesi are more closely related to each other, forming a clade with slightly weaker support that is sister to C. moluccensis from Aru Island, Maluku. Similarly, C. granularis shares morphological similarities with C. pseudorugosus, as supported by the molecular phylogeny that places them as sister species. However, although C. perlatus and C. purpureus exhibit morphological similarities to these two species, their relationships are more distant (Fig. 23), suggesting that these shared traits may have evolved convergently. Coenobita moluccensis, C. patsyae, and C. celebensis exhibit varying degree of yellow or orange coloration on the cheliped meri, and even on the second and third pereopods (Figs. 4, 8, 12). In C. lila, a very faint yellow tinge is also present on the pereopods (Rahayu et al. 2016: figs. 10, 11A–C). According to the phylogenetic tree (Fig. 23), the presence of yellow or orange coloration on the cheliped meri and other pereopods is likely a derived trait in these closely related species. Some individuals of C. patsyae have brightly yellow or orange cheliped meri (Fig. 8D), which are very similar to those in C. celebensis (Fig. 12), suggesting a closely relationship between them, as supported by the phylogenetic tree and haplotype network (Figs. 23, 24). Species discovery and geographic distribution In our study, four new species and one newly recorded species of Coenobita are reported from Indonesia, increasing the total number of recognized species in this country to 13: C. brevimanus, C. cavipes, C. celebensis, C. granularis, C. lila, C. longitarsis, C. moluccensis, C. patsyae, C. perlatus, C. pseudorugosus, C. rugosus, C. variabilis, and C. violascens. Consequently, Indonesia stands out as the global hotspot with the highest species diversity for this genus, a pattern similar to that observed in other marine organisms (Veron et al. 2009; Siallagan et al. 2023). Notably, six of these species, viz. C. celebensis, C. granularis, C. patsyae, C. pseudorugosus, C. rugosus, and C. violascens, can be found within the Gulf of Tomini, Central Sulawesi (Shih et al. 2023a; this study), suggesting that unique habitat conditions in this area may contribute to speciation within the genus. It is recommended that further studies on larval ecology (e.g., Hamasaki et al. 2015 2018; Doi et al. 2018), population ecology (e.g., Barnes 2002 2003; Bundhitwongrut et al. Table 3. Matrix of the percentage of pairwise nucleotide divergences with Kimura 2-parameter (K2P) distances and the number of basepair (bp) differences based on the cytochrome c oxidase subunit I (COI) gene within and between the new species (and new record) of Coenobita in this study and their closely related species. In the interspecific (right) part of the table, lower-left values represent K2P distances and upper-right values indicate bp differences. Range of values are given in parentheses Intraspecific Interspecific Nucleotide divergence Mean nucleotide difference C. patsyae C. celebensis C. moluccensis C. lila C. variabilis C. granularis C. pseudorugosus C. patsyae 1.22 (0–3.3) 7.86 (0–21) 30.38 (23–35) 33.03 (23–39) 66.74 (61–70) 65.66 (60–69) 91.66 (84–97) 93.38 (87–97) C. celebensis 0.86 (0.15–1.39) 5.6 (1–9) 4.81 (3.6–5.58) 31.23 (23–35) 65 (63–67) 71.88 (67–75) 94.66 (89–100) 94.42 (91–98) C. moluccensis 1.14 (0–2.02) 7.36 (0–13) 5.27 (3.61–6.29) 4.96 (3.61–5.6) 65.25 (60–69) 60.92 (57–65) 94.76 (88–100) 92.91 (88–96) C. lila 1.08 7 11.21 (10.14–11.83) 10.87 (10.5–11.23) 10.96 (9.99–11.67) 78.57 (78–79) 91.7 (87–95) 85.5 (83–88) C. variabilis 0.98 (0–1.86) 6.38 (0–12) 10.99 (9.94–11.62) 12.12 (11.2–12.72) 10.12 (9.4–10.87) 13.41 (13.3–13.49) 84.85 (78–90) 85.5 (84–87) C. granularis 1.35 (0–2.33) 8.77 (0–15) 15.67 (14.17–16.74) 16.25 (15.13–17.33) 16.33 (14.98–17.39) 15.7 (14.77–16.37) 14.37 (13.05–15.37) 76.64 (72–83) C. pseudorugosus 0.64 (0.46–0.92) 4.17 (3–6) 16.03 (14.77–16.74) 16.22 (15.54–16.93) 15.97 (14.99–16.61) 14.43 (13.96–14.92) 14.47 (14.17–14.77) 12.9 (12.02–14.13) page 35 of 38Zoological Studies 64:11 (2025) © 2025 Academia Sinica, Taiwan 2014), and population genetics (e.g., Hamasaki et al. 2017), as these areas of research may help to elucidate the mechanisms of sympatry and provide valuable insights for conservation efforts. Considering that Coenobita species from other regions of Indonesia have not been well studied, we anticipate that more species may be discovered in the future. CONCLUSIONS In our study, we confirmed the existence of four new species, viz. C. moluccensis n. sp., C. patsyae n. sp., C. celebensis n. sp., and C. granularis n. sp., as well as one newly recorded species, C. variabilis, in Indonesia. This confirmation is based on the integration of morphological characters and molecular evidence derived from mitochondrial 16S and COI markers. Notably, C. moluccensis, C. patsyae, and C. celebensis, supported by genetic evidence, form a major clade that can be considered as a species complex. This complex is closely related to C. lila, with which it shares morphological similarities. Additionally, C. granularis and C. pseudorugosus exhibit similar morphological traits and are identified as sister species in our analysis. The discovery of C. variabilis in West Papua, Indonesia, expands its known distribution from Australia to New Guinea Island. Our research increases the total number of Coenobita species documented in Indonesia to 13. This highlights the effectiveness of integrative taxonomy in confirming the existence of distinct species and contributes to our understanding of the biodiversity of Coenobita in Indonesia. Moreover, it underscores the importance of considering multiple sources of data in species identification and classification. Acknowledgments: This study was supported by a grant from the National Science and Technology Council (NSTC 112-2313-B-005-051-MY3), Executive Yuan, Taiwan, to HTS. We thank Min-Wan Chen for helping part of the molecular work; Peter Davie and Marissa McNamara for loaning specimens in QM; and Choi Sin Tung (Tony Choi) for providing useful information. We also acknowledge the two anonymous referees who helped improve the manuscript. Authors’ contributions: HTS conceived this study, morphological description, molecular analysis, and drafted the manuscript. DLR performed the morphological description, line drawings, and drafted the manuscript. FAP provided the specimens, performed the ecological observation and color in life, and drafted the manuscript. All authors read and approved the final manuscript. Competing interests: The authors declare that they have no conflict of interest. Availability of data and materials: Sequences generated in the study were deposited into the GenBank database (accession numbers in Table 1). Consent for publication: Not applicable. Ethics approval consent to participate: Not applicable. REFERENCES Alcock A. 1905. Catalogue of the Indian Decapod Crustacea in the Collection of the Indian Museum. Part II. Anomura. Fasciculus I. Pagurides. Trustees of the Indian Museum, Calcutta, India, xi+193 pp., 16 pls. Barnes DKA. 2002. 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