Pagurion tuberculata Shiino, 1933

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 : 112-118

Pagurion tuberculata Shiino, 1933

( Figs. 18 , 19 )

Pagurion tuberculata Shiino, 1933: 254–256 , fig. 2 [Tanabe Bay, Japan; infesting Pagurus watasei Terao, 1913 = Dardanus aspersus (Berthold, 1845) ]; Shiino, 1952: 41 [mention]; Danforth, 1963: 9 [list]; Shiino, 1972: 7 [list]; Harada, 1991: 201 [list]; Saito et al., 2000: 36 [list]; Markham, 2003: 72 [list]; Madad, 2008: 2, 5, 6, 17, 33–34, 46, 48, 51, 86, 87, fig. 9 [Batangas, Philippines; infesting Dardanus lagopodes (Forskål, 1775) ]; Yu & An, 2008: 692 [list]; Markham, 2010: 151, 152, 156–158, figs. 6, 7 [Queensland, Australia; infesting Dardanus arrosor (Herbst, 1796) , D. hessii (Miers, 1884) ]; McDermott et al., 2010: 8 [list]; An et al., 2013: 561–563, fig. 1 [South China Sea; infesting D. aspersus ]; An et al., 2016: 45 [list]; Williams & Boyko, 2016: 36 [list].

Pagurion tuberculatum [sic] Dollfus in Neveu-Lemaire et al., 1934: 74 [list].

Pagurion arrosor An, Li & Markham, 2013: 561 , 563–564, fig. 2 [ South China Sea, Stn. 6238, 20°00′N, 108°00′E, 83 m; infesting D. arrosor ]; new synonymy.

Material examined. Philippines: 1 juvenile ♀ (1.4 mm) ( USNM 1494005 ) from right branchial chamber of juvenile ♂ Dardanus sp. (1.3 mm SL), Batangas, Anilao, coll. JDW, 13 February 1999 ; 1 dextral ♀ (7.7 mm), 1 mature ♂ (3.00 mm) ( USNM 1494006 , ♂ on SEM stub) from right branchial chamber of ♀ Dardanus lagopodes (4.5 mm SL), Batangas, Sombrero Island , coll. JDW, 10 June 2000 . Indonesia: 1 ovigerous sinistral ♀ (13.7 mm), 1 mature ♂ (3.6 mm, lost) ( ZRC 2018.0829 ), from left branchial chamber of Dardanus megistos (Herbst, 1804) (10.2 mm SL), East Lombok , Ekas, coll. 23 July 2009 .

Type locality. Tanabe Bay , Japan ( Shiino, 1933) .

Distribution. Tanabe Bay, Japan ( Shiino, 1933); Batangas, Philippines ( Madad, 2008; herein), South China Sea (An et al., 2013); Queensland, Australia (Markham, 2010).

Hosts. Diogenidae . Dardanus arrosor (Herbst, 1796) (Markham, 2010; An et al., 2013); D. aspersus (Berthold, 1845) ( Shiino, 1933; An et al., 2013); D. hessii (Miers, 1884) (Markham, 2010) ; D. lagopodes (Forskål, 1775) ( Madad, 2008; herein), D. megistos (Herbst, 1804) (herein).

Size range (length). Females to 14.0 mm, males to 4.5 mm ( Shiino, 1933).

Remarks. The mature females of Pagurion tuberculata examined ( Fig. 18A, B ) do not differ from the original type material as described by Shiino (1933) or the subsequent records of Markham (2010) and An et al. (2013). The female has five pairs of biramous pleopods, the barbula has a series of lateral projections, the inner ridge of oostegite 1 is digitate and the uropods are biramous, which are all consistent with the diagnosis provided by Shiino (1933). The Philippine male of P. tuberculata ( Figs. 18C , 19A ) exhibits the same article counts for the antennae, 3 antennular and 4 antennal ( Fig. 19B, C ), as described by Shiino (1933). An et al. (2013), however, illustrated 3 antennular and 5 antennal articles for this species. The original description of P. tuberculata did not include a detailed description of the pereopods of males ( Fig. 19D, E ) but their character states (first two pairs larger than others, though not markedly so) was mentioned by Markham (2010) and An et al. (2013). Markham (2010) stated that his female specimens had 2 pairs of coxal plates, not 4 as in the original description, but from his illustrations it appears that he mixed up the counts for dorsolateral bosses and coxal plates because his illustrated specimen has 2 dorsolateral bosses on each side and 4 coxal plates. Males have 5 pairs of broad pleopods ( Fig. 19F ); Markham (2010) noted that his male had 5 pairs of pleopods and Shiino (1933) stated there were also 5 pairs but that the posterior 2 pairs were “rather inconspicuous.”

The genus Pagurion Shiino, 1933 , monotypic until recently, is composed of P. tuberculata and P. arrosor An, Li & Markham, 2013 . Genus-defining characteristics of the females include having the cephalon separated from the thorax, distinct thoracic segments, rudimentary coxal plates present on the first four segments, oostegites forming a complete marsupium with no medial gaps, distinct abdominal segments ( P. tuberculata with a distinctly visible pleotelson, P. arrosor with an obscure pleotelson), lamellar lateral plates in all six segments, and five pairs of biramous pleopods as well as biramous uropods ( Shiino, 1933; An et al., 2013). Males of Pagurion exhibit a head separated from the thorax, distinct thoracic and abdominal segments (although one specimen of P. tuberculata was reported with fusion between the last two pleomeres, as was the allotype of P. arrosor ; see Markham, 2010; An et al., 2013), and uniramous tuberculate pleopods and no uropods ( Shiino, 1933).

The status of P. arrosor as a distinct species is not supported by the evidence. An et al. (2013) stated that the differences between P. arrosor and P. tuberculata were: females lacking frontal laminae (present in P. tuberculata ), antennae “1 and 2” (antennules and antennae) of 4 and 5 articles each, respectively (3 and 4 articles in P. tuberculata ), pleomere 6 (pleotelson) “obscure” = dorsally not visible (the female has 6 pleomeres, but the pleotelson is covered by the fifth pleomere lateral plates) (“distinct” = dorsally visible in P. tuberculata ), males with last two pleomeres somewhat fused (separate in P. tuberculata ). However, most of these purported differences do not hold up under scrutiny. Pagurion arrosor was illustrated by An et al. (2013) with a small but present frontal lamina, contradicting the text description. An et al. (2013) also reported a female of P. tuberculata with 3 antennular and 5 antennal articles, not the 3 and 4 articles they indicate to be the character state for P. tuberculata in their comparison with P. arrosor , indicating that the antennal counts are variable in the species. Markham (2010) illustrated a female of P. tuberculata from Dardanus arrosor as having both pleomeres 5 and 6 “obscure” (i.e., obscured by the fourth pleomere) and the male having the last two pleomeres partially fused. These characters were used by An et al. (2013) to define, in part, P. arrosor ; however, they included Markham’s (2010) specimens as a valid record of P. tuberculata rather than P. arrosor without explanation. An et al. (2013) stated in the abstract that characters of the barbula and pleopods of the female also distinguished P. arrosor from P. tuberculata , but they did not state what these differences were, nor did they mention them in the body of the text. We can see no differences in the pleopod morphology of the females, based on the descriptions and illustrations of Shiino (1933), Markham (2010) and An et al. (2013), as well as the present material. The barbulae of the specimens of P. tuberculata reported by An et al. (2013) have little to no digitation in the medial regions, but the barbulae of the syntype female of P. tuberculata and the holotype of P. arrosor are virtually identical. An et al. (2013) reported the male of P. arrosor to have midventral tubercles on the pereomeres; these are also present in our specimens but were not previously reported from P. tuberculata males. However, they are small and not easily observed, and may well have been overlooked by Shiino (1933) and others. Based on the above comparison and clear indication of intraspecific variability in some characters of both males and females, we consider P. arrosor to be a synonym of P. tuberculata .

Markham (2010) gave the type host as “ Dardanus sp.” but the host was identified to species in Shiino (1933), although it was at that time usually considered a synonym of D. scutellatus (H. Milne Edwards, 1848) until later synonymised with D. aspersus by Osawa (2013). An et al. (2013) were apparently unaware of Osawa’s (2013) paper and stated that their record from D. aspersus was the first of any bopyrid on that host.

The juvenile hermit crab host reported here (USNM 1494005) An J, Markham JC & Yu H (2010) Description of two new species appears to also be a Dardanus but this is not certain and and a new genus of bopyrid isopod parasites (Bopyridae: it is not identifiable to species. Markham (2010) neither Pseudioninae) of hermit crabs from China. Journal of Natural described nor illustrated his specimens of P. tuberculata History, 44: 2065–2073.

An J, Yu H & Williams JD (2007) Four new records and a new from D. hessi (Miers, 1884), so it is not known what the species of Dactylokepon Stebbing, 1910 (Epicaridea: Bopyridae: character states were for those specimens; An et al. (2013) Ioninae) from Chinese waters. Journal of Natural History, 41: did not include this host record in their synonymy list. An 2063–2079.

et al. (2013) incorrectly cited Calcinus gaimardii (H. Milne An J, Zhao Q & Markham JC (2016) Paguristione uniuropodus, a Edwards, 1848) and Calcinus minutus Buitendijk, 1937 as new genus and a new species of Pseudioninae infesting hermit hosts of P. tuberculata and incorrectly credited these host crabs from China (Crustacea, Isopoda, Bopyridae). ZooKeys, records as coming from Madad (2008); no such data appears 577: 43–53.

in that thesis. An et al. (2013) also incorrectly ascribed the Anderson G & Dale WE (1981) Probopyrus pandalicola (Packard) erroneous combination “ Clibanarius gaimardii ” (= Calcinus (Isopoda, Epicaridea): morphology and development of larvae in culture. Crustaceana, 41: 143–161.

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Figures

Fig. 18. Pagurion tuberculata Shiino, 1933, mature female, USNM 1494006 (A, B), juvenile female, USNM 1494005 (C). A, female, dorsal view; B, female, ventral view; C, juvenile female, dorsal view. Scale bars = 1 mm (A, B); 250µm (C).

Fig. 19. Pagurion tuberculata Shiino, 1933, male, USNM 1494006 (A–F). A, ventral overview; B, right antennule; C, right antenna; D, right pereopods 1 and 2; E, right pereopod 7; F, pleopods. Scale bars = 500 µm (A, F); 50 µm (B, C); 200 µm (D); 100 µm (E).