Bopyrissa marami, Williams & Boyko & Madad, 2019

Williams, Jason D., Boyko, Christopher B. & Madad, Asma Z., 2019, Branchial parasitic isopods (Crustacea: Isopoda: Bopyridae: Pseudioninae) of hermit crabs (Crustacea: Decapoda: Diogenidae) from the western Pacific, with descriptions of a new genus and three new species, Raffles Bulletin of Zoology 67, pp. 83-118 : 96-104

Bopyrissa marami , new species

(Figs. 7–12)

Pseudione sp. A Madad, 2008: 2 , 4–6, 37–45, 48, 49, 64, 88–99, figs. 10–15, table 3 [ Philippines; infesting Calcinus gaimardii (H. Milne Edwards, 1848) , C. latens (Randall, 1840) , and C. minutus Buitendijk, 1937 ].

Material examined. Philippines: Dextral ♀ holotype (4.2 mm) ( USNM 1493918 ), mature ♂ allotype (1.5 mm) ( USNM 1493919 ), from right branchial chamber of ♂ Calcinus gaimardii (2.9 mm SL), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; Paratypes: 1 dextral ovigerous ♀ (4.0 mm), 1 ♂ (1.6 mm) ( USNM 1493920 , on SEM stubs) from right branchial chamber of ♀ C. gaimardii (3.2 mm SL), Cebu, Olango Island, coll. JDW, 9 July 1997 ; 1 dextral ♀ (4.9 mm) hyperparasitised by Cabirops sp. , 1 ♂ (1.6 mm) (JDW pers. coll.) from right branchial chamber of ♂ C. gaimardii (3.9 mm SL), Batangas, Sombrero Island, coll. JDW, 13 July 1997 ; 1 dextral ovigerous ♀ (3.8 mm) ( USNM 1493921 ) from right branchial chamber of ♀ Calcinus latens (Randall) (3.0 mm SL), Batangas, Sombrero Island, coll. JDW, 13 July 1997 ; 1 dextral ovigerous ♀ (4.4 mm), 1 ♂ (1.50 mm) ( USNM 1493922 ) from right branchial chamber of ♂ C. gaimardii (2.9 mm SL), Batangas, Sombrero Island, coll. JDW, 13 July 1997 ; 1 dextral ovigerous ♀ (5.0 mm), 1 ♂ (1.7 mm) ( USNM 1493923 ) from right branchial chamber of ♂ C. gaimardii (3.6 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 10 July 1997 ; 1 dextral ovigerous ♀ (4.7 mm), 1 ♂ (1.8 mm) ( USNM 1493924 ) from right branchial chamber of ♀ C. gaimardii (3.6 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 10 July 1997 ; 1 dextral ovigerous ♀ (4.2 mm), 1 ♂ (1.7 mm) ( USNM 1493925 ) from right branchial chamber of ♂ C. gaimardii (3.9 mm SL), Puerto Galera , Big Lalaguna Beach, coll. JDW, 31 July 1997 ; 1 dextral ovigerous ♀ (4.2 mm), 1 ♂ (1.5 mm) ( USNM 1493926 ) from right branchial chamber of ♀ C. gaimardii (3.3 mm SL), Puerto Galera , Big Lalaguna Beach, coll. JDW, 31 July 1997 ; 1 dextral ovigerous ♀ (4.3 mm), 1 ♂ (1.3 mm) ( USNM 1493927 ) from right branchial chamber of ♀ C. gaimardii (3.2 mm SL), Puerto Galera , Big Lalaguna Beach, coll. JDW, 31 July 1997 ; 1 dextral ovigerous ♀ (3.9 mm), 1 ♂ (1.50 mm) ( USNM 1493928 ) from right branchial chamber of ovigerous ♀ C. gaimardii (3.5 mm SL), Puerto Galera , Big Lalaguna Beach, coll. JDW, 31 July 1997 ; 1 dextral ovigerous ♀ (3.16 mm), 1 ♂ (1.06 mm) ( USNM 1493929 ), from right branchial chamber of ♀ C. gaimardii (2.3 mm SL), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ovigerous ♀ (3.5 mm), 1 ♂ (1.4 mm) ( USNM 1493930 ), from right branchial chamber of ♀ C. gaimardii (2.2 mm SL), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ♀ (2.9 mm), 1 ♂ (1.1 mm) ( USNM 1493931 ), from right branchial chamber of ♂ C. gaimardii (2.4 mm SL), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ♀ (5.0 mm), 1 ♂ (1.9 mm) ( USNM 1493932 ), from right branchial chamber of ♂ C. gaimardii (3.3 mm SL), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ovigerous ♀ (4.4 mm), 1 ♂ (1.4 mm) ( USNM 1493933 ), from right branchial chamber of ♂ C. gaimardii (3.1 mm SL), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ovigerous ♀ (3.66 mm), 1 ♂ (1.30 mm), from right branchial chamber of ♂ Calcinus minutus (2.7 mm SL), host also parasitised by an entoniscid (JDW pers. coll.), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ovigerous ♀ (3.7 mm), 1 ♂ (1.3 mm) ( USNM 1493934 ), from right branchial chamber of ♂ C. gaimardii (2.9 mm SL), host also infested with Eremitione clibanaricola , new combination ( USNM 1493999 ), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ovigerous ♀ (4.3 mm), 1 ♂ (1.4 mm) ( USNM 1493935 ), from right branchial chamber of ♂ C. minutus (3.2 mm SL), Puerto Galera , Coco Beach, coll. JDW, 12 January 1999 ; 1 dextral ovigerous ♀ (4.6 mm), 1 ♂ (1.6 mm) ( USNM 1493936 ), from right branchial chamber of ♂ C. gaimardii (3.3 mm SL), Batangas, Sombrero Island, coll. JDW, 30 January 1999 ; 1 dextral ♀ (3.4 mm), 1 ♂ (1.1 mm) ( USNM 1493937 ), from right branchial chamber of ♂ C. gaimardii (2.5 mm SL), Batangas, Sombrero Island, coll. JDW, 30 January 1999 ; 1 dextral ovigerous ♀ (3.9 mm), 1 ♂ (1.4 mm) ( USNM 1493938 ) from right branchial chamber of ♂ C. gaimardii (3.1 mm SL), Batangas, Anilao, coll. JDW, 13 February 1999 ; 1 dextral ovigerous ♀ (4.3 mm), 1 ♂ (1.3 mm) ( USNM 1493939 ), from right branchial chamber of ♀ C. gaimardii (3.0 mm SL), Batangas, Anilao, coll. JDW, 13 February 1999 ; 1 dextral ovigerous ♀ (5.5 mm), 1 ♂ (1.7 mm) ( USNM 1493940 ), from right branchial chamber of ♂ C. gaimardii (3.8 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (3.4 mm), 1 ♂ (1.3 mm) ( USNM 1493941 ), from right branchial chamber of ♂ C. gaimardii (2.7 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (4.6 mm), 1 ♂ (1.2 mm) ( USNM 1493942 ), from right branchial chamber of ♀ C. gaimardii (3.2 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (5.9 mm), 1 ♂ (1.6 mm) ( USNM 1493943 ), from right branchial chamber of ♂ C. gaimardii (4.9 mm SL), host also parasitised by a peltogastrid rhizocephalan (host and parasite sent to R. Yoshida), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (3.0 mm), 1 ♂ (1.1 mm) ( USNM 1493944 ), from right branchial chamber of ♀ C. gaimardii (2.1 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (2.60 mm), 1 ♂ (1.19 mm) ( USNM 1493945 ), from right branchial chamber of ♂ C. gaimardii (2.5 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (2.9 mm), 1 ♂ (1.2 mm) ( USNM 1493946 ), from right branchial chamber of ♂ C. gaimardii (2.5 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (5.2 mm), 1 ♂ (1.8 mm) (JDW pers. coll.), from right branchial chamber of ♀ C. gaimardii (3.2 mm SL), host also parasitised by a peltogastrid rhizocephalan, Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (4.6 mm), 1 ♂ (1.5 mm) ( USNM 1493947 ), from right branchial chamber of ♀ C. gaimardii (3.2 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (3.6 mm), 1 ♂ (1.2 mm) ( USNM 1493948 ), from right branchial chamber of ♂ C. gaimardii (3.0 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (not measured, dried), 1 ♂ (not measured, dried) (JDW pers. coll.), from right branchial chamber of ♀ C. gaimardii (2.1 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral mature ♀ (4.7 mm), 1 ♂ (1.5 mm) ( USNM 1493949 ), from right branchial chamber of ♀ C. gaimardii (3.9 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (4.5 mm), 1 ♂ (1.2 mm) ( USNM 1493950 ), from right branchial chamber of ♀ C. gaimardii (2.9 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (5.2 mm), 1 ♂ (1.9 mm) ( USNM 1493951 ), from right branchial chamber of ♀ C. gaimardii (3.6 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (3.0 mm), 1 ♂ (1.2 mm) ( USNM 1493952 ), from right branchial chamber of ♂ C. gaimardii (2.6 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (2.6 mm), 1 ♂ (1.1 mm) ( USNM 1493953 ), from right branchial chamber of ♂ C. gaimardii (2.3 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral mature ♀ (4.8 mm), 1 ♂ (1.5 mm) ( USNM 1493954 ), from right branchial chamber of ♀ C. gaimardii (3.0 mm SL), Bataan, Mabayo, coll. JDW, 21 February 1999 ; 1 dextral ovigerous ♀ (4.6 mm), 1 ♂ (1.5 mm) ( USNM 1493955 ), from right branchial chamber of ♀ C. gaimardii (3.6 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral ovigerous ♀ (3.72 mm), 1 ♂ (1.25 mm) ( USNM 1493956 ), from right branchial chamber of ♂ C. gaimardii (2.6 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral ♀ (5.6 mm), 1 ♂ (1.3 mm) ( USNM 1493957 ; male on SEM stub, JDW pers. coll.), from right branchial chamber of ♂ C. gaimardii (3.4 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral juvenile ♀ (1.4 mm), 1 ♂ (0.6 mm) (JDW pers. coll.), from right branchial chamber of ♀ C. gaimardii (1.8 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral mature ♀ (5.1 mm), 1 ♂ (1.8 mm) ( USNM 1493958 ), from right branchial chamber of ♀ C. gaimardii (3.4 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral juvenile ♀ (2.8 mm), 1 ♂ (1.0 mm) ( USNM 1493959 ), from right branchial chamber of ♀ C. gaimardii (2.3 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral ovigerous ♀ (4.1 mm), 1 ♂ (1.3 mm) ( USNM 1493960 ), from right branchial chamber of ♀ C. gaimardii (2.7 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral ovigerous ♀ (5.3 mm), 1 ♂ (1.7 mm) ( USNM 1493961 ), from right branchial chamber of ♀ C. gaimardii (3.5 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral mature ♀ (3.0 mm), 1 ♂ (1.1 mm) ( USNM 1493962 ), from right branchial chamber of ♀ C. gaimardii (2.4 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral ovigerous ♀ (3.1 mm), 1 ♂ (1.1 mm) ( USNM 1493963 ), from right branchial chamber of ♀ C. gaimardii (2.62 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral ovigerous ♀ (4.0 mm), 1 ♂ (1.4 mm) ( USNM 1493964 ), from right branchial chamber of ♀ C. gaimardii (2.6 mm SL), Bataan, Morong, coll. JDW, 28 February 1999 ; 1 dextral ovigerous ♀ (2.5 mm), 1 ♂ (1.0 mm) ( USNM 1493965 ), from right branchial chamber of ♂ C. gaimardii (1.9 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 27 March 1999 ; 1 dextral ovigerous ♀ (4.2 mm), 1 ♂ (1.5 mm), 1 cryptoniscus larva of unidentified Cabirops sp. (0.8 mm) (JDW pers. coll.), from right branchial chamber of ♀ C. gaimardii (3.4 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 27 March 1999 ; 1 dextral ovigerous ♀ (4.0 mm), 1 ♂ (1.4 mm) ( USNM 1493966 ), from right branchial chamber of ♀ C. gaimardii (2.7 mm SL), Boracay , White Beach, coll. JDW, 15 April 1999 ; 1 dextral ovigerous ♀ (5.5 mm), 1 ♂ (1.6 mm) ( USNM 1493967 ), from right branchial chamber of ♂ C. gaimardii (3.8 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ovigerous ♀ (4.4 mm), 1 ♂ (1.3 mm) ( USNM 1493968 ), from right branchial chamber of ♂ C. gaimardii (3.1 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ♀ (2.7 mm), 1 ♂ (1.0 mm) ( USNM 1493969 ), from right branchial chamber of ♂ C. gaimardii (2.3 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ovigerous ♀ (5.6 mm), 1 ♂ (1.8 mm) ( USNM 1493970 ), from right branchial chamber of ♂ C. gaimardii (4.2 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ovigerous ♀ (4.3 mm), 1 ♂ (1.3 mm) ( USNM 1493971 ), from right branchial chamber of ♂ C. gaimardii (3.2 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ovigerous ♀ (5.0 mm), 1 ♂ (1.4 mm) ( USNM 1493972 ), from right branchial chamber of ♂ C. gaimardii (3.7 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ovigerous ♀ (6.1 mm), 1 ♂ (2.0 mm) ( USNM 1493973 ), from right branchial chamber of ♂ C. gaimardii (4.4 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ovigerous ♀ (6.2 mm), 1 ♂ (1.6 mm) ( USNM 1493974 ), from right branchial chamber of ♂ C. gaimardii (4.3 mm SL), Bataan, Morong, coll. JDW, 25 April 1999 ; 1 dextral ovigerous ♀ (5.3 mm), ♂ (1.6 mm) ( USNM 1493975 ), from right branchial chamber of ♂ C. gaimardii (3.5 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (4.0 mm), 1 ♂ (1.2 mm) ( USNM 1493976 ), from right branchial chamber of ♂ C. gaimardii (2.7 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (5.1 mm), 1 ♂ (1.6 mm) ( USNM 1493977 ), from right branchial chamber of ♂ C. gaimardii (3.5 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (5.0 mm), 1 ♂ (2.1 mm) ( USNM 1493978 ), from right branchial chamber of ♂ C. gaimardii (3.8 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral mature ♀ (4.7 mm), 1 ♂ (1.8 mm) ( USNM 1493979 ), from right branchial chamber of ♀ C. gaimardii (3.5 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (3.8 mm), 1 ♂ (1.3 mm) ( USNM 1493980 ), from right branchial chamber of ♂ C. gaimardii (2.7 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (4.5 mm), 1 ♂ (1.4 mm) ( USNM 1493981 ), from right branchial chamber of ♂ C. gaimardii (3.2 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (3.9 mm), 1 ♂ (1.2 mm) ( USNM 1493982 ), from right branchial chamber of ♀ C. gaimardii (2.9 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (3.5 mm), 1 ♂ (1.0 mm) ( USNM 1493983 ), from right branchial chamber of ♂ C. gaimardii (2.7 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (3.9 mm), 1 ♂ (1.3 mm) ( USNM 1493984 ), from right branchial chamber of ♂ C. gaimardii (2.8 mm SL), Batangas, Sombrero Island, coll. JDW, 10 June 2000 ; 1 dextral ovigerous ♀ (3.7 mm), 1 ♂ (1.3 mm) ( USNM 1493985 , on SEM stub), from right branchial chamber of ♀ C. gaimardii (3.3 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 18 June 2000 ; 1 dextral ovigerous ♀ (5.6 mm), 1 ♂ (1.8 mm) ( ZRC 2018.0818 ), from right branchial chamber of ♂ C. gaimardii (3.9 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 18 June 2000 ; 1 mature ♀ (4.5 mm) ( ZRC 2018.0820 ), from right branchial chamber of ♀ C. gaimardii (3.1 mm SL), host also parasitised by an entoniscid (host in JDW pers. coll.), Puerto Galera , Lalaguna Beach, coll. JDW, 18 June 2000 ; 1 mature ♀ (4.3 mm), 1 ♂ (1.53 mm) ( ZRC 2018.0821 ), from right branchial chamber of ovigerous ♀ C. minutus (3.2 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 18 June 2000 ; 1 ovigerous ♀ (4.9 mm), 1 ♂ (1.8 mm) ( ZRC 2018.0823 ), from right branchial chamber of ♀ C. gaimardii (3.2 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 18 June 2000 ; 1 mature ♀ (3.9 mm), 1 ♂ (1.4 mm) ( ZRC 2018.0825 ), from right branchial chamber of ♂ C. minutus (2.9 mm SL), Puerto Galera , Lalaguna Beach, coll. JDW, 18 June 2000 .

Description. Female body length 4.10 mm, maximal width 2.41 mm, head length 0.66 mm, head width 1.01 mm, pleon length, 0.9 mm. Mean length of female specimens was 4.39 ± 0.93 mm and mean maximal width was 2.57 ± 0.52 mm (n=48). Head dextrally deflexed with ~40° distortion angle. Body pyriform, all body regions and segments distinct (Figs. 7A, B, 8A, B). Head suboval, deeply set into pereon; frontal lamina semicircular, notched slightly in middle, covered in scales (Figs. 7B, 8A); eyes absent. Antennules of 3 articles each, article 1 of right and left antennules fused; setae at distal tip of antennules and scattered on ventral surface and margins of article 1; scales covering entire surface (Figs. 8C, 9B). Antennae of 5 articles each, covered in scales, tuft of setae at distal tip and setae on lateral surface and at margins of articles (Figs. 8C, 9A). Maxilliped with acute spur, slender palp with ~7 setae present, posterior margin of maxilliped setose (Figs. 8L, M, 9C, D). Barbula with two lateral projections with variable digitation, medial projection most variable, sometimes as a low rounded nub (Fig. 8D–I). Oostegites completely enclosing brood chamber; setae on posterior margin of fifth oostegite; oostegite 1 with rounded anterior lobe, triangular posterior lobe with short setae along edge, inner ridge of oostegite 1 with 6–7 large digitate projections (Figs. 8J, K, 9E, F). Pereon of 7 distinct pereomeres, broadest across pereomere 4, tapering towards pleon posteriorly. Prominent dorsolateral bosses and coxal plates on segments 1–4, tergal projections half length of dorsolateral bosses on segments 1–4; segments 5–7 with dorsolateral bosses continuing whole length of segment, tergal projections absent (Figs. 7A, 8A). Pereopods 1–5 subequal in size, 6 and 7 slightly smaller, setae on anterior tip of propodi, directly underneath dactyli, no large gap between pereopods, scales on all pereopodal articles (Figs. 8N, O, 9G). Pleon of 6 distinct pleomeres, sixth pleomere ventrally displaced, first 5 pleonal segments with lateral plates. Five pairs of biramous pleopods, tuberculate on margins; small pair of lanceolate, distally rounded, uniramous uropods present on pleomere 6, uropods mostly hidden in dorsal view (Figs. 7B, C, 8A, 9H).

Male length 1.62 mm, maximal width 0.59 mm at pereomere 4, head length 0.13 mm, head width 0.31 mm, pleon length 0.41 mm. Mean length of male specimens was 1.55± 0.24 mm and mean maximal width was 0.58 ± 0.08 mm (n=48). Head subovate, distinct from pereomere 1, all segments distinct (Figs. 10A, 11A); prominent eyes on posteriolateral border of head (Fig. 10A). Antennules of 2 articles each, setae on distal tip of article 2, setae on surface of article 2; antennae of 2 articles each, tuft of setae on distal tip of antennae and along ventral surface of article 2, scales on surface of article 2 (Figs. 10C, 11B). Pereon of 7 distinct pereomeres, broadest across pereomere 4, tapering posteriorly. Pereopods 3 and 4 largest, size decreasing anteriorly and posteriorly with pereopods 6 and 7 smallest, ventral surface of carpi and meri covered in scales, propodi with ridges of scales on anterior portion underneath dactyl, tuft of setae on anterior tip of propodi opposed to tip of dactyli (Figs. 10D, E, 11C, D). Pleon of 5 distinct pleomeres, tapering posteriorly, distolateral edge fringed with setae (Figs. 10A, 11E). Pleopods tuberculate, present on pleomeres 1–4 (Figs. 10B, 11E). Pleotelson with weak anterolateral projections, distolateral ends with setae; anal fissure slightly produced ventrally, midventral tubercles absent, no uropods (Figs. 10B, F–H, 11E, F).

Type locality. Coco Beach , Puerto Galera, Philippines .

Etymology. The species epithet is derived from the Filipino word marami , which means “many” or “numerous,” reflecting the fact that this was by far the most common bopyrid found on hermit crabs from the Philippines during this study.

Distribution. Batangas, Puerto Galera, Philippines.

Hosts. Diogenidae : Calcinus gaimardii (H. Milne Edwards, 1848) , C. latens (Randall, 1840) , and C. minutus Buitendijk, 1937 .

Size range (length). Females to 6.2 mm, males to 2.1 mm.

Remarks. Descriptions and diagnoses of species of Pseudione and closely related genera are often problematic, given the intraspecific variability and inconsistency of supposedly uniform features ( Markham, 1978). The new species described herein is superficially similar to Eremitione clibanaricola , new combination, but it does not exhibit the marked asymmetry in female specimens that is characteristic of E. clibanaricola , new combination, nor does E. clibanaricola , new combination present a ventrally displaced sixth pleomere, which is characteristic of all Bopyrissa species. Females of B. marami , new species are most similar to those of Bopyrissa guamensis An, Gong, and Paulay, 2018 , recently described from Calcinus elegans (H. Milne Edwards, 1836) collected in Guam. However, females of B. marami , new species are distinguished from B. guamensis based primarily on characters of the first oostegite, in which the posterior lobe is recurved and with smaller and more numerous projections on the inner ridge in B. guamensis but rounded and with fewer, larger projections on the inner ridge in B. marami , new species. The pleopods of the males of B. marami , new species are different from those of B. guamensis in that the former species has 4 pairs of medially situated pleopods whereas the latter has 3 pairs (lacking the pair on the fourth pleomere). The new species is also similar to B. liberorum Markham, 1985b , but differences between females of the two species include: two lateral lobes on the barbula in B.

H

Fig. 10. Bopyrissa marami , new species, male allotype, USNM 1493919 (A–E) and paratypes, USNM 1493975 (F), USNM 1493933 (G),

USNM 1493970 (H). A, dorsal view; B, ventral view; C, right antennule and antenna; D, right pereopod 1; E, left pereopod 7; F–H, male pleons. Scale bars = 100 µm (A, B, F–H); 20 µm (C); 50 µm (D, E).

marami , new species vs. one lobe in B. liberorum ; inner ridge of oostegite 1 with approximately 8 large digitiform extensions in B. marami , new species vs. 2 or 3 low lobes in B. liberorum ; maxilliped with extended palp in B. marami , new species vs. maxilliped with recessed palp in B. liberorum ; pleopods with tubercles in B. marami , new species vs. smooth in B. liberorum . Differences between males of the two species include: pleopods large and positioned medially in B. marami , new species vs. pleopods low and positioned laterally in B. liberorum ; posterolateral extentions of the pleotelson weakly extended in B. marami , new species vs. strongly produced in B. liberorum .

The position of the uniramous uropods of females of B. marami , new species is much like that of B. novaeguineensis ( Danforth, 1971) , new combination; however, there is no setose palp on the maxilliped of females of B. novaeguineensis , new combination. Males of B. novaeguineensis , new combination have low laterally placed pleopods and the overall body shape is much more elongate than for any specimens of B. marami , new species.

Ecology. Prevalence of Bopyrissa marami , new species over the thirteen collections of hermit crabs in which the parasite was found ranged from 0.8–19.2% (mean prevalence=6.4±6.4%, n=1,671 total hermit crabs examined).

A ².5

)

mm ².0

(

Length 1.5

Total 1.0

Male 0.5

Of these specimens, the vast majority (91.7%) infested Calcinus gaimardii but the species was also found on one C. latens and five C. minutus ( Table 1). Two specimens of B. marami , new species were hyperparasitised by cabiropid isopods (2.6%, 2 of 78). One host C. gaimardii was doubly infested with Eremitione clibanaricola , new combination and B. marami , new species and two C. gaimardii were doubly parasitised by a peltogastrid rhizocephalan and B. marami , new species. Two hosts (one C. gaimardii and one C. minutus ) were doubly parasitised by an unidentified entoniscid and B. marami , new species.

The mean total body length of female B. marami , new species was 4.2 ± 0.9 (n=77); male B. marami , new species mean total body length was 1.4 ± 0.3 (n=75). The relationship between female total body length and male total body length was significantly positively correlated (P <0.0001) (Fig. 12A). Bopyrissa marami , new species was found to be ovigerous in all months sampled (January through April, June, and July) and thus likely breeds throughout the year, as does its dominant host C. gaimardii ( Litulo & Tudge, 2005) . Of the 75 mature female B. marami , new species collected, 84% were ovigerous; the smallest ovigerous female was 2.5 mm SL.

The total body length of female B. marami , new species and host hermit crabs (as estimated by carapace length) was significantly positively correlated (P <0.0001) (Fig. 12B). One female C. gaimardii host (USNM 1493928) was found to be ovigerous although infested with a mature female and male pair of B. marami , new species; none of the other female hosts were found with eggs.

Variation. The number of articles and characteristics of the antennules of female specimens of Bopyrissa marami , new species showed some variation; specifically, the specimen prepared for SEM exhibited two articles, with article 1 of the left and right antennules being fused. The antennules of this specimen were shorter and more setose over the entire surface of article 1 than in other specimens ( Fig. 9A, B ). The barbula was typically composed of one or two digitate projections, the size of the projections and amount of digitation was variable (Fig. 8D–I); only two female specimens (4.2%; two of 48) exhibited a barbula represented by a single, slender projection, lacking digitation, similar to that of E. clibanaricola , new combination (USNM 1493945, USNM 1493974). The degree of digitation of the inner ridge of oostegite 1 was variable, with some female specimens showing weaker digitation than others but with digitation always present to some degree. A maxilliped palp was present in all females, with some specimens having a much larger, more pronounced setose palp than others. The length of the uropods varied; in some specimens, they were longer than the lateral plates while in other specimens, the uropods were considerably shorter. One female specimen exhibited biramous uropods, possibly the result of a developmental abnormality (USNM 1493969); a few other female specimens exhibited some degree of abnormality such as an undeveloped pleomere 1 on one side and coalescing of one or more pleomeres. Male morphology was fairly consistent overall, although they varied in the characteristics of the pleotelson, which could be almost cordiform in morphology or slightly cylindrical (Fig. 10A, F–H).

Figures

Fig. 9. Bopyrissa marami, new species, female paratype, USNM 1493920. A, left antenna; B, Left antennule, terminal article; C, left maxilliped, outer view; D, palp of maxilliped; E, right oostegite 1, internal view; F, right oostegite 1, digitate inner ridge; G, left pereopod 1; H, pleopods, lateral view. Scale bars = 200 µm (A, F, G); 50 µm (B, D); 500 µm (C, E, H).