taxonID	type	description	language	source
03B6E97BFFC8CA4DFF61ABD75894F81E.taxon	type_taxon	Type genus: Diogenion Codreanu, Codreanu & Pike, 1960	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC8CA4DFF61ABD75894F81E.taxon	diagnosis	Diagnosis: As for genus, see below. Included genera: Diogenion Codreanu, Codreanu & Pike, 1960 (1 species).	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC8CA4DFF61ABD75894F81E.taxon	discussion	Remarks: As discussed by Adkison (1990), Diogenion exhibits two apomorphic characters among entoniscids: females with seven pairs of pereopods and lack of pleopods. This genus appears to be the most primitive group of entoniscids and sister taxon to the other two subfamilies, Entoniscinae (apomorphies: females with host derived sheath and oostegites forming brood pouch, first oostegite not different from others, males with unsegmented pereopods) and Entionine (apomorphies: females with heart in pleomere 3 and at least some pleural lamellae complexly folded). Specimens belonging to Diogenion contain characters found in both previously recognized subfamilies; specifically, they share characters with species in Entioninae (presence of a maxilliped in females, and males with segmented pereopods) and Entoniscinae (heart in pleomere 1, absence of pleural lamellae and uropods in females). In terms of larval development, the sole species of Diogenion exhibits synchronous development as is found in species of Entioninae, in contrast to the asynchronous development (i. e., multiple stages at one time in brood chamber) in species of Entoniscinae. Although Adkison (1990) concluded that Diogenion should belong in its own subfamily, he referred to the taxon only as “ Subfamily A ” throughout the work and his dissertation was never published. In addition, Adkison (1990) never examined any specimens of Diogenion; his conclusions were made solely from analysis of the original description (Codreanu et al. 1960). Based on our examination of new material of Diogenion cf. vermifactus from the Philippines and confirmation of the unique suite of characters, we conclude that the genus should be placed in its own subfamily as designated herein.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC9CA4CFF61AFB25F31FDDA.taxon	diagnosis	Diagnosis: Female head with dorsal groove; antennulae broad, antennae reduced; small maxilliped present. Oostegites 1 – 7 present; oostegite 1 without ascendant lobe, with strongly developed recurrent lobe lined with pleatlike folds; brood pouch formed by overlapping of oostegites 1 – 4, oostegites 5 – 7 reduced. Pereopods 1 – 7 present. Pleon slender, heart tubercle in pleomere 1; pleural lamellae, pleopods and uropods absent. Male with both pairs of antennae present, antennulae projecting beyond margin of head; head not fused with pereomere 1. Pereopods 1 – 6 present, multisegmented; lacking pereopod 7 or greatly reduced. Pleon of five segments plus pleotelson, medioventral spines absent, uropods present. Epicaridium larva with pereopods 1 – 6 isomorphic; lacking pereopod 7; pereopod 6 similar in length to pereopod 5. Pleon with five pairs of biramous pleopods. Pleotelson tapering to point. Uropods with endopod slightly longer than exopod. Cryptoniscus larva unknown.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC9CA40FF61AD4E5FFDFEDE.taxon	description	Figures 3 – 5	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC9CA40FF61AD4E5FFDFEDE.taxon	materials_examined	Material examined. Philippines: Female (19.4 mm) (USNM 1522331), infesting Calcinus gaimardii (3.5 mm SL), inhabiting unknown shell, Lalaguna Beach, Puerto Galera, 13 ° 31 ’ 32 ″ N, 120 ° 58 ’ 8 ″ E, coll. J. D. Williams, 3 March 1999. Female (6.9 mm), male (2.5 mm), and multiple larvae on two SEM stubs (USNM 1522332), infesting Calcinus pulcher (3.5 mm SL), inhabiting unknown shell, Batangas, Anilao, 13 ° 42 ’ 14.5 ″ N, 120 ° 52 ’ 45.3 ″ E, coll. J. D. Williams, 13 February 1999. Female (19.0 mm) (USNM 1522333), infesting C. gaimardii (6.0 mm SL), inhabiting unknown shell, Mabayo, Bataan, 14 ° 44 ′ 00 ″ N, 120 ° 16 ′ 32 ″ E, coll. J. D. Williams, 21 February 1999. Female (10.5 mm) and male (2.5 mm) (USNM 1522334), infesting Calcinus minutus (2.7 mm SL), inhabiting unknown shell, host also with bopyrid parasite (Bopyrissa marami) in right branchial chamber, Coco Beach, Puerto Galera, 13 ° 31 ′ 32 ″ N, 120 ° 57 ′ 44 ″ E, coll. J. D. Williams, 14 January 1999 (indicated as 12 January 1999 in Williams et al. 2019). Female (15.2 mm) and male (no measurement) (ZRC 2022.0962), infesting C. gaimardii (4.4 mm SL), inhabiting unknown shell, Lalaguna Beach, Puerto Galera, 13 ° 31 ′ 32 ″ N, 120 ° 58 ′ 8 ″ E, coll. J. D. Williams, 17 June 2000. Female (4.6 mm) and male (1.5 mm) (ZRC 2016.0411) from male Pagurojacquesia polymorpha (2.3 mm), inhabiting unknown shell, west off Panglao Island, Station CP 2334, 09 ° 37.5 ′ 00 ″ N, 123 ° 40.2 ′ 00 ″ E (see de Forges et al., 2009), beam trawl, 631 – 659 m, sandy bottom, Panglao Expedition 2005, 22 May 2005.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC9CA40FF61AD4E5FFDFEDE.taxon	biology_ecology	Hosts: Calcinus gaimardii, Calcinus minutus, Calcinus pulcher, Diogenes senex (type host of Diogenion vermifactus), Pagurojacquesia polymorpha.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC9CA40FF61AD4E5FFDFEDE.taxon	description	Description of female: Female in abdomen of host surrounded by host induced sheath; positioned with anterior end at posterior end of host and ventral surface of parasite directed to dorsal surface of host (fig. 3 A, B). Host tube with exit pore near first pereopod of host (fig. 3 B). Head bilobed, with elongated oval lobes (figs. 3 C, 4 A). Antennulae fused into large, flattened, pad-like structure. Antennae as thin digitate extensions (fig. 4 A). Maxillipeds small, round (fig. 4 A). Pereon without ovarian processes. Oostegites 1 – 7 present, progressively smaller posteriorly; oostegite 1 normally folded back posterior to head and surrounded by other oostegites (fig. 3 A, B), large and flap-like with rounded anterior end and inner surface lined with pleat-like folds (fig. 3 C). Pereopods 1 – 7 present; pereopods segmented, each with small dactylus present on distal end; scales present (fig. 4 B, C). Pleon slender, with five pleomeres; small heart in pleomere 1; pleopods, pleural lamellae and uropods absent (fig. 4 D). Brood development synchronous; all larvae at same stage of development. Description of male: Male found within brood chamber of female. Only one male accompanying female (when present). Body curved ventrally, pale in color with few spots of pigmentation (fig. 4 E, F). Head small and rounded. Pair of elongated conical antennulae extending beyond margin of head, terminally setose; pair of conical antennae slightly larger than antennulae each with tuft of terminal setae (fig. 4 G). Oral cone with mandibles and maxilliped present (fig. 4 G). Seven pereomeres; pereon maximal width at pereomeres 4 – 6, gradually tapering anteriorly and posteriorly (fig. 4 E, F). Six pairs of pereopods present, absent on pereomere 7 (fig. 4 F); pereopods multi-segmented, subequal in size, each with curved dactylus (fig. 4 H); patch of scales on propodus at point where dactylus terminates on all pereopods (fig. 4 H). Pleon of five segments plus pleotelson; pleomeres gradually decreasing in size posteriorly; without appendages (fig. 4 E, F). Uropods well developed; terminal setae present (fig. 4 E). Description of epicaridium larva (undergoing molt): Approximately 340 µm in length. Body teardrop shaped and dorsally convex; anterior margin of head rounded (fig. 5 A). Head with pair of conical antennulae (segmentation and setae not distinct due to molting process) (fig. 5 A, C); pair of long antennae of seven segments, four peduncular and three flagellar, half of total body length when fully extended (fig. 5 A, B). Oral cone with pair of maxillipeds on each side of mouth on ventral side. Pereon of seven segments. Six pereopods; subequal in length, subchelate with ovate propodi and curved dactyli (fig. 5 A, B). Pleon with five pairs of biramous pleopods, subequal in size, with three short setae on exopod and two short setae on endopod (fig. 5 D). Telson tapered to point. Uropods with pointed endopod, slightly longer than exopod, both bearing short terminal setae (fig. 5 E).	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC9CA40FF61AD4E5FFDFEDE.taxon	discussion	Remarks: All specimens match the original morphological description of Diogenion vermifactus (Codreanu et al. 1960), previously known only infesting Diogenes senex in the Red Sea (fig. 2). However, in this study D. cf. vermifactus was found to parasitize three species of shallow subtidal calcinid hermit crabs, C. gaimardii, C. minutus and C. pulcher and the deeper water pagurid P. polymorpha in the coastal waters of the Philippines, over 8,000 km away from the type locality in the Red Sea (fig. 1). Currently, we cannot morphologically distinguish the Philippine specimens of Diogenion from those of the Red Sea; however, given this disjunct distribution and the barriered nature of the Red Sea (DiBattista et al. 2016; Schnurr et al. 2018; Hadfield & Smit 2020) it is possible that the Philippine entoniscid is a distinct species from D. vermifactus. Additional data, ideally including molecular sequencing, is needed to clarify whether D. vermifactus is widely distributed across the Indo-West Pacific or represents a species complex (Hadfield & Smit 2020). At present, the substantial geographic distance between the two localities and diversity of hosts suggests that a designation of “ cf. ” for the Philippine specimens is warranted pending discovery of additional specimens from along the range. Many features of the adult Diogenion vermifactus are considered primitive when compared to other members of Entoniscidae (Codreanu et al. 1960; Adkison 1990). Specifically, the female has segmented pereopods, a brood pouch formed by overlapping oostegites and a synchronously developing brood; males also retain segmented pereopods and possess both pairs of antennae as well as uropods. For these reasons, Adkison (1990) suggested that the genus be placed in its own subfamily. The epicaridium larval stage of D. vermifactus was not previously described except for mention of the narrowness of the larvae, the great length of the appendages and the uniform structure of the pereopods and pleopods (Codreanu et al. 1960; Adkison 1990). Although previous authors did not provide drawings or detailed descriptions of this larval stage, the present specimens exhibit the gross morphological details reported previously. The present study confirms that the epicaridium larvae have nearly uniform pereopods, i. e., the sixth pereopod is not greatly modified as in the larvae of some entoniscid genera (e. g., Cancrion Giard & Bonnier, 1887). However, the epicaridium larvae examined were undergoing a molt, making some features difficult to distinguish, as has been shown to occur in other epicarideans (see Williams & Boyko 2021).	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC5CA40FF61AE935F1AFCFA.taxon	diagnosis	Diagnosis: Female head with dorsal groove; antennulae and antennae present as fused pads; maxilliped present. Pereon without ovarian processes; oostegites 1 – 5 present; oostegite 1 largest with ascendant and recurrent lobes; other oostegites on posterior of pereon, directed dorsally; oostegite 5 reduced. Heart in pleomere 1; five pairs of pleopods present; posterior margin of pleomere 6 entire; uropods absent. Male antenna projecting beyond margin of head; pereopods unsegmented; medioventral spines and pleopods absent; posterior margin of pleomere 6 entire. Epicaridium larva pereopod 6 similar in length to pereopod 5, with reduced dactylus; pleopods 1 – 3 with medial angle of peduncle bearing one long seta, exopod bearing three long setae; pleopod 4 with medial angle of peduncle bearing one short seta, exopod bearing three long setae; pleopod 5 reduced, setae absent. Cryptoniscus larva unknown. Included species: Paguritherium alatum Reinhard, 1945; P. manggagaway n. sp.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC5CA42FF61ACAF58F9FC9E.taxon	description	Figure 6	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC5CA42FF61ACAF58F9FC9E.taxon	materials_examined	Material examined: United States: Ovigerous female (8.5 mm), mature male (0.9 mm) (USNM 1522335), infesting male Pagurus longicarpus (2.7 mm) inhabiting unknown shell, inside Hereford Inlet on the Cape May peninsula of New Jersey, 39 ° 02 ′ N, 74 ° 48 ′ W, coll. J. J. McDermott, 10 July 1985. Ovigerous female (5.6 mm) (USNM 1522336), infesting female P. longicarpus (2.5 mm) inhabiting unknown shell, inside Hereford Inlet on the Cape May peninsula of New Jersey, 39 ° 02 ′ N, 74 ° 48 ′ W, coll. J. J. McDermott, 18 June 1986. Ovigerous female (9.5 mm) with eggs and few epicaridium larvae on SEM stub (USNM 1522337), infesting female P. longicarpus (2.6 mm) inhabiting unknown shell, inside Hereford Inlet on the Cape May peninsula of New Jersey, 39 ° 02 ′ N, 74 ° 48 ′ W, coll. J. J. McDermott, 12 August 1987. Epicaridium larva (304 µm; JDW pers. coll. 9 - 12 - 17), infesting Acartia hudsonica Pinhey, 1926 (1.3 mm), taken in plankton tow at town of Hempstead East Marina, Point Lookout, New York. 40 ° 35 ′ 37.70 ″ N, 73 ° 35 ′ 6.39 ″ W, coll. J. D. Williams, 12 September 2017.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC5CA42FF61ACAF58F9FC9E.taxon	description	Brief remarks on male morphology (USNM 1522336): The males (fig. 6 E – H) have six unsegmented pereopods, reduced to rounded knob-like structures, covered in spinous scales (fig. 6 E, G, H). The “ spines ” reported by Reinhard (1945; fig. 6 F herein) are actually scales as are found in a wide range of epicarideans (e. g., Nielsen & Strömberg 1969; Cericola & Williams 2015). Pereopod 7 is very reduced, with only a minute, scale-covered lobe present as shown with SEM (fig. 6 G). Brief remarks on epicaridium larval morphology (USNM 1522337): Adkison (1990) was the first to report on the morphology of the epicaridium larvae of P. alatum (see fig. 6 I – N), showing that they have: antennulae of three segments; antennae of six segments; pereopods 1 – 5 gnathopodal, without specialized fan-like setae; pereopod 6 with a reduced triangular dactylus; pleopod 5 setae absent, exopod reduced; and uropods biramous with the exopod longer than the endopod. Adkison (1990) mislabeled pereopod 6 as pereopod 7 for the epicaridium larvae of this species in his figure 10 (fig. 10 h should be labeled as pereopod 6; fig. 10 F and G are more anterior pereopods, but it cannot be determined which ones; these are reproduced and labeled as such herein fig. 6 K – M). Aside from this error, the morphology of the epicaridium larvae described by Adkison (1990) is confirmed based on the new material herein studied, including the lack of specialized fan-like setae on pereopods 1 – 5 and pereopod 6 with a reduced triangular dactylus (fig. 6 N).	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC5CA42FF61ACAF58F9FC9E.taxon	discussion	Remarks: The adult specimens of P. alatum examined here were those reported in McDermott (1998) from New Jersey where he found four out of 3,703 (0.11 %) P. longicarpus infested. The female specimens match the original description (fig. 6) and the redescription of the species in Adkison′s (1990) unpublished dissertation which was based on the syntypes (USNM 81563) and four additional specimens (USNM 235935). McDermott (1998) found that the pleon of females extended to an opening toward the base of the eyestalks. Adkison (1990) indicated that in at least one specimen the pleon extended to an opening toward the base of the maxilliped. During sampling of plankton on Long Island, New York, one copepod (Acartia hudsonica) was found with an epicaridium larva attached that matched in all aspects the morphology indicated above for P. alatum. This is the first confirmed identification of any entoniscid from a copepod intermediate host in nature (see notes in Discussion on experimental settlement of entoniscid epicaridium larvae on copepods).	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC7CA58FF61AC8B58A2FA26.taxon	description	Figures 7 – 11	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC7CA58FF61AC8B58A2FA26.taxon	materials_examined	Material examined: Philippines. Ovigerous female holotype (9.9 mm; USNM 1522338), ovigerous female paratype (8.1 mm; USNM 1522339), male allotype (0.8 mm; USNM 1522340 from paratype female), infesting male Calcinus gaimardii (3.6 mm SL), inhabiting unknown shell, Lalaguna Beach, Puerto Galera, 13 ° 31 ′ 32 ″ N, 120 ° 58 ′ 8 ″ E (type locality), coll. J. D. Williams, 31 July 1997. Ovigerous female paratype (10.5 mm) (USNM 1522341), infesting male C. gaimardii (5.5 mm SL), inhabiting unknown shell, Lalaguna Beach, Puerto Galera, 13 ° 31 ′ 32 ″ N, 120 ° 58 ′ 8 ″ E, coll. J. D. Williams, 28 July 1997. Two ovigerous female paratypes (9.0 mm and 7.5 mm) (USNM 1522342), infesting male C. gaimardii (4.1 mm SL), inhabiting unknown shell; ovigerous female paratype (15.9 mm) (USNM 1522343), infesting female C. latens (3.0 mm SL), inhabiting unknown shell, Lalaguna Beach, Puerto Galera, 13 ° 31 ′ 32 ″ N, 120 ° 58 ′ 8 ″ E, coll. J. D. Williams, 31 July 1997. Ovigerous female paratype (9.8 mm) (USNM 1522344), infesting female Calcinus latens (5.2 mm SL), inhabiting unknown shell, Sombrero Island, Puerto Galera, 13 ° 41 ′ 53 ″ N, 120 ° 49 ′ 47 ″ E, coll. J. D. Williams, 5 July 1997; ovigerous female paratype (6.5 mm) (USNM 1522345), infesting female C. latens (3.7 mm SL), inhabiting unknown shell, Sombrero Island, Puerto Galera, 13 ° 41 ′ 53 ″ N, 120 ° 49 ′ 47 ″ E, coll. J. D. Williams, 5 July 1997. Female paratype (7.5 mm) and multiple larvae on SEM stub (USNM 1522340), infesting male C. gaimardii (5.4 mm SL), inhabiting unknown shell, Lalaguna Beach, Puerto Galera, 13 ° 31 ′ 32 ″ N, 120 ° 58 ′ 08 ″ E, coll. J. D. Williams, 3 March 1999. Female paratype (broken, not measured; USNM 1522347), infesting male C. gaimardii (4.0 mm SL), inhabiting unknown shell, Batangas, Anilao, 13 ° 42 ′ 14.5 ″ N, 120 ° 52 ′ 45.3 ″ E, coll. J. D. Williams, 13 February 1999. Female paratype (broken, not measured) and male paratype (2.4 mm) (ZRC 2022.0963), infesting female C. gaimardii (4.2 mm SL), inhabiting unknown shell, Batangas, Anilao, 13 ° 42 ′ 14.5 ″ N, 120 ° 52 ′ 45.3 ″ E, coll. J. D. Williams, 13 February 1999. Female (8.5 mm) and male (0.8 mm) (ZRC 2022.0964), infesting unidentified hermit crab (damaged), Sombrero Island, Puerto Galera, 13 ° 41 ′ 53 ″ N, 120 ° 49 ′ 47 ″ E, coll. J. D. Williams, 10 June 2000. Female paratype (8.0 mm) and multiple larvae on SEM stub (ZRC 2022.0965), female infesting C. gaimardii (3.1 mm SL), inhabiting Drupella cornus (Röding, 1798), host also with bopyrid parasite in right branchial chamber (Bopyrissa marami; ZRC 2018.0820), Lalaguna Beach, Puerto Galera, 13 ° 31 ′ 32 ″ N, 120 ° 58 ′ 08 ″ E, coll. J. D. Williams, 18 June 2000.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC7CA58FF61AC8B58A2FA26.taxon	etymology	Etymology: The specific name is from the Tagalog deity Manggagaway, one of the goddesses of Kasamaan (underworld) opposed to Bathala (the supreme being, maker of all things) and blamed as the cause of disease (Piscos 2019).	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC7CA58FF61AC8B58A2FA26.taxon	description	Description of female (based on holotype, USNM 1522338, and paratypes): Female in abdomen of host surrounded by host-induced sheath. Female positioned with anterior end at posterior end of host and ventral surface of parasite directed to dorsal surface of host (fig. 7 A). Pleon extended into thoracic cavity of host and continuing to exit pore in middle of fifth pereomere (fig. 7 A), or more anteriorly, including to branchial chamber, mouthparts, or at base of host eye stalk. Head bilobed (Figs. 8, 9 A, B), with approximately spherical lobes. Antennules and antennae as fused pads anterior to head (fig. 9 A, B). Pair of small maxillipeds present. Five pairs of oostegites present; oostegite 1 largest, ascendant lobe extending anterior to head and smaller recurrent lobe (Figs. 7 B – E, 8 A). Pleon slender, stalk-like, of five pleomeres plus pleotelson (fig. 9 C, D); pleomeres subequal in size, tapering slightly posteriorly; channel formed between five pairs of long pleopods, at least last two pairs of pleopods extending beyond pleotelson (fig. 9 C, D). Uropods absent. Rounded heart in pleomere 1 (Figs. 9 D). Brood development asynchronous; multiple stages of development observed in single female (Figs. 7, 8 A). Description of male (based on allotype, USNM 1522340, and paratypes): Male found within brood chamber. In present samples, only one male accompanying female (when present). Body curved ventrally, pale in color with few spots of pigmentation (fig. 9 E). Short blunt head fused with pereomere 1; antennulae large, lobelike, extending posterolaterally with 16 – 18 setae. Oral cone large; rounded maxillipeds present (fig. 9 G). Pereon of seven pereomeres, six pairs of small rounded pereopods on pereomeres 1 – 6; increasing slightly in size from pereopod 1 to 5, sixth pereopod subequal in size to fourth; pereopods unsegmented and distally covered in scales (fig. 9 E, H), pereopods absent or each reduced to small nodule on pereomere 7 (fig. 9 E). Pleon of five segments plus pleotelson; pleomeres gradually smaller in size posteriorly, without appendages, pleotelson undivided; uropods absent (fig. 9 E). Description of epicaridium larva (based on USNM 1522340 and ZRC 2022.0965): Approximately 373 µm in length (fig. 10 A). Body teardrop shaped, dorsally convex; anterior margin of head rounded (fig. 10 A, B). Oral cone with pair of maxillipeds present on ventral side (fig. 10 C). Antennulae of three short, rounded segments subequal in length; basal segments wider than terminal segment; terminal segment with three long setae and several short setae (fig. 10 C, D). Antenna of six segments, four peduncular and two flagellar; three short setae and one longer stout seta at end of first flagellar segment, one long serrated seta with annulation at base at end of second flagellar segment; antenna approximately half total body length when fully extended (fig. 10 A, D). Pair of pereopods present on pereomeres 1 – 6; five anterior pairs equal in size (fig. 10 A, B); pereopods 1 – 5 subchelate, each with ovate propodus with few short setae and long curved dactylus; dactylus approximately half length of propodus (fig. 10 E, F); one broad serrated fan-like seta and one more elongate denticulate seta on propodus opposite dactylus of first five pereopods (fig. 10 F). Pereopod 6 of four segments, slightly shorter than preceding pereopods; merus short, carpus and propodus fused, lanceolate in shape with few short setae and scales; dactylus reduced to small triangular extension (fig. 11 A – C). Pleon with five pairs of biramous pleopods, pleopods 1 – 4 subequal in size, with three long setae on truncate exopod and one long setae on pointed endopod (fig. 11 A, D); pleopod 5 reduced, without setae (fig. 11 E). Uropods biramous; exopod slightly larger than endopod, with one long seta and one short seta, endopod with one long seta (fig. 11 F).	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
03B6E97BFFC7CA58FF61AC8B58A2FA26.taxon	discussion	Remarks: Paguritherium was previously a monotypic genus, with P. alatum as the type species (Reinhard 1945; Adkison 1990). Paguritherium manggagaway n. sp. matches the diagnostic features of the genus (e. g., female body is slender and lacking ovarian processes; four pairs of well-developed oostegites, fifth pair rudimentary; male antennulae tuberculiform and pereopods stump-like) (Reinhard 1945). The females of P. manggagaway n. sp. bear a strong resemblance to those of P. alatum but differ in the length of the pleopods (P. manggagaway n. sp. has much longer pleopods relative to body length than P. alatum). Ovigerous females of P. manggagaway n. sp. could be detected in fixed host hermit crab specimens prior to dissection in some cases. The head of the entoniscid was sometimes visible through the host abdominal cuticle and the abdomen of these hosts was more distended than those of uninfested individuals; in addition, the eggs and larvae of the entoniscid were visible through the host abdominal cuticle in some cases (fig. 7). Reinhard (1945) and Adkison (1990) also indicated that eggs and larvae of P. alatum could be seen through the host abdomen. No developing juvenile females (stages 3 – 9 sensu Kuris et al. 1980) of P. manggagaway n. sp. were found, even in those host specimens systematically dissected; these stages may be relatively short in duration. Reinhard (1945) also found no juvenile females in his samples of P. alatum. As in P. alatum, the pleon of P. manggagaway n. sp. can extend to an exit pore near the anterior end of the host, including the host eyestalks, but can also be found terminating near the mouthparts, branchial chamber or pereomeres. Although the pleon of most female entoniscids end in the branchial chamber, several species of entoniscids have a larval exit pore on or around the host eye stalk (Shiino 1942; Adkison 1990). In one of the doubly infested hermit crabs, the pleon of the posterior female paratype extended to the base of the pleon of the anterior female holotype (fig. 7 B – E); it appears that the host derived sheath connected the two females, such that the epicaridium larvae of both females would be released through the exit pore of the more anterior female. Two female paratypes (USNM 1522341 and 1522347) were found with calcified parts (fig. 8 B) and the bodies in poor condition. This could be due to host response to parasitism (Kuris et al. 1980). Other studies have noted the presence of dead female entoniscids encapsulated or degenerating within host crabs (Miyashita 1941; Shiino 1942) but did not comment further upon parasite mortality. In some cases, the entoniscids can be rendered nearly unrecognizable due to encapsulation and calcification by the host (Kuris et al. 1980). The male of P. manggagaway n. sp. is very similar to that of P. alatum, but with more pronounced antennulae. Regarding the epicaridium larvae, P. manggagaway n. sp. can be distinguished from P. alatum based on the presence of specialized fan-like seta on the propodus of each of the first five pereopods. This feature was not found in P. alatum by Adkison (1990) or in the present samples of that species. Although it is possible that they were overlooked, these setae are relatively large and have been documented in other entoniscids. For example, such setae were described from Entoniscus japonicus Shiino, 1942 infesting the anomuran host Petrolisthes japonicus (De Haan, 1849) native to the Indo-West Pacific (Shiino 1942). These setae were described as curved, forked spines or serrated disks on the margin facing the dactylus on the propodus of each of the first five pereopods (Shiino 1942). The annulation of the terminal setae on the antennae (see Watling 1989) of P. manggagaway n. sp. has not been reported before in entoniscids, but it may have been previously overlooked in studies which did not use SEM. Unfortunately, the description of entoniscid larvae (epicaridium, microniscus and cryptoniscus) is lacking for many species, requiring new collections of entoniscids and ideally examination of these stages with SEM. The annulation of the distal seta makes it appear to be a third flagellar segment, but this is unlikely because nearly all other entoniscid epicaridium larvae have four peduncular and two flagellar antennal segments. There are likely other differences in the antennulae, antennae, and mouth parts of the epicaridium larvae between the two species of Paguritherium; future analyses using SEM should be completed on epicaridium and cryptoniscus larvae of P. alatum from the east coast of the United States and P. manggagaway n. sp.	en	Detorre, Marissa, Williams, Jason D., Boyko, Christopher B. (2023): A review of the endoparasitic isopods (Epicaridea: Entoniscidae) from hermit crabs, including description of the new subfamily Diogenioninae and a new species of Paguritherium Reinhard, 1945 from the Philippines. Zootaxa 5249 (1): 12-40, DOI: 10.11646/zootaxa.5249.1.2, URL: http://dx.doi.org/10.11646/zootaxa.5249.1.2
