Research Article |
Corresponding author: Aoi Tsuyuki ( tykamsp0430@gmail.com ) Academic editor: Tom Artois
© 2020 Aoi Tsuyuki, Hiroshi Kajihara.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Tsuyuki A, Kajihara H (2020) A giant new species of Enchiridium (Polycladida, Prosthiostomidae) from southwestern Japan. ZooKeys 918: 15-28. https://doi.org/10.3897/zookeys.918.47061
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We describe a new species of polyclad flatworm, Enchiridium daidai sp. nov., from the rocky subtidal zone in the East China Sea along the coasts of the Kyushu and Okinawa Islands, Japan. Enchiridium daidai sp. nov. is characterized by i) the entire periphery of the dorsal surface narrowly fringed with orange, ii) a marginal-eyespot band extending to the position of the mouth (about anterior one-eighth of body), and iii) two prostatic vesicles covered by a common muscle sheath, which is penetrated by the ejaculatory duct. We performed a molecular phylogenetic analysis based on 945-bp 28S rDNA sequences of 16 species of Prosthiostomidae currently available in public databases in addition to those of E. daidai sp. nov. and Prosthiostomum torquatum Tsuyuki et al., 2019. In the resulting tree, our new species was nested in a clade composed of Enchiridium species. The tree topology was in favor of a taxonomic view that Enchiridium should be defined by having i) a common muscle sheath that encloses two prostatic vesicles and ii) marginal eyespots that may or may not surround the periphery of the dorsal surface.
Cotylea, marine flatworm, phylogeny, Platyhelminthes, taxonomy
Polyclad flatworms in the family Prosthiostomidae Lang, 1884 are characterized by i) an elongated body with a ventral sucker after the female gonopore, ii) a plicate tubular pharynx, and iii) paired prostatic ducts, each of which extends from a spherical prostatic vesicle and enters the penis or the ejaculatory duct independently, instead of uniting to each other before the entrance. Prosthiostomidae is composed of five genera: Enchiridium Bock, 1913; Enterogonimus Hallez, 1913; Euprosthiostomum Bock, 1925; Lurymare Du Bois-Reymond Marcus & Marcus, 1968; and Prosthiostomum Quatrefages, 1845 (
In Japan, 21 species of prosthiostomids were previously reported, but there was no record of Enchiridium (
Three polyclad specimens were collected subtidally from under rocks in Kagoshima and Okinawa, southwestern Japan (Fig.
For histological examination, tissues were dehydrated in an ethanol series, cleared in xylene, embedded in paraffin wax, and sectioned at a thickness of 7 µm using a microtome. Sections were stained with hematoxylin and eosin, mounted on glass slides in Entellan New (Merck, Germany), and then observed and photographed under an Olympus BX51 compound microscope. All type slides have been deposited in the Invertebrate Collection of the Hokkaido University Museum (ICHUM), Sapporo, Japan.
Total DNA was extracted using a silica-based method (
Additional 28S rDNA sequences were downloaded from GenBank; Pseudobiceros splendidus (Lang, 1884) (Pseudocerotidae) and Prostheceraeus crozieri (Hyman, 1939) (Euryleptidae) were chosen as outgroups (Table
All graphical treatments were done with Adobe Photoshop CC. Illustrations were prepared with Adobe Illustrator CC.
List of species used for the molecular phylogenetic analysis and respective GenBank accession numbers.
Species | GenBank accession number | Reference |
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Prosthiostomidae | ||
Enchiridium daidai sp. nov. | LC504235 | This study |
LC504236 | ||
LC504237 | ||
Enchiridium evelinae Marcus, 1949 | KY263683 |
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Enchiridium japonicum Kato, 1943 | MH700298 |
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Enchiridium periommatum Bock, 1913 | MH700299 |
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MH700300 | ||
MH700301 | ||
Enchiridium sp. 1 | MH700302 |
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MH700303 | ||
Enchiridium sp. 2 | MN384686 |
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Enchiridium sp. 3 | KY263673 |
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Enchiridium sp. 4 | KY263679 |
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Euprosthiostomum mortenseni Marcus, 1948 | MH700304 |
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Prosthiostomum acroporae (Rawlinson et al., 2011) | HQ659011 |
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Prosthiostomum cynarium Marcus, 1950 | MH700371 |
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Prosthiostomum lobatum Pearse, 1938 | MH700372 |
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Prosthiostomum milcum Du Bois-Reymond Marcus & Marcus, 1968 | MH700373 |
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Prosthiostomum purum Kato, 1937 | MH700374 |
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Prosthiostomum siphunculus (Delle Chiaje, 1822) | HQ659012 |
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Prosthiostomum torquatum Tsuyuki et al., 2019 | LC504234 | This study |
Prosthiostomum trilineatum Yeri & Kaburaki, 1920 | MH700376 |
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Prosthiostomum utarum Marcus, 1952 | MH700377 |
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Prosthiostomum sp. | MH700375 |
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Outgroup | ||
Prostheceraeus crozieri (Hyman, 1939) | HQ659013 |
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Pseudobiceros splendidus (Lang, 1884) | MH700388 |
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Genus Enchiridium Bock, 1913 sensu
The new specific name daidai is a Japanese noun, meaning the color orange. It was named after the thin marginal orange line surrounding the entire dorsal fringe.
Three specimens, all collected by A. Tsuyuki. Holotype:
Off the coast of Bonomisaki (31.2542N, 130.2150E), Kagoshima, Japan.
Body elongated, tapered posteriorly, 28–77 mm long (77 mm in holotype) and 4.6–14 mm maximum width (14 mm in holotype) in living state (Fig.
Subtidal (5–14 m depth), under rocks.
Specimens from Kagoshima and Okinawa differed in body size. The holotype from Kagoshima was 77 mm long and 15 mm wide, whereas the paratype specimens from Okinawa were 28–37 mm long and 4.6–7.4 mm wide (Fig.
Body elongated, usually rounded anteriorly; dorsal surface translucent, fringed by a thin marginal orange line; marginal eyespots present only anteriorly; plicated pharynx tubular in shape, about one-fifth of body length; pair of prostatic vesicles bound by common muscular sheath, the latter penetrated by ejaculatory duct.
In the phylogenetic tree, Enchiridium daidai sp. nov. was nested in a clade composed of Enchiridium species (Fig.
In spite of the noticeable difference in body size, specimens from Kagoshima and Okinawa – all having reached sexual maturity – were identified as conspecific. They shared the following morphological characteristics: i) body dorsally fringed with a thin orange line, ii) marginal-eyespot band extending to the position of the mouth (about anterior one-eighth of the body), iii) two prostatic vesicles covered by a common muscle sheath, and iv) common muscle sheath penetrated by ejaculatory duct. In addition, the COI p-distances among the specimens, 0.002–0.012, fell in a range of intraspecific values, 0.000–0.020, which was observed in four species of the acotylean leptoplanoid Notocomplana (
As for the taxon concept of Enchiridium, our results did not show a compatibility to
Enchiridium daidai sp. nov. is distinguished from E. evelinae, E. japonicum, E. periommatum, and E. punctatum by the arrangement of the marginal eyespots; the marginal-eyespot band in these four species completely encircles the periphery of the dorsal surface, whereas that of our specimens is present only along the anterior margin. Our new species is also easily distinguished from the other four congeners, E. delicatum, E. gabriellae, E. magec, and E. russoi, by the thin marginal orange line surrounding the entire dorsal fringe and by the lack of spots or maculae on the dorsal surface (Table
Reaching 77 mm in body length, Enchiridium daidai sp. nov. is the largest species in the genus over E. punctatum (about 40 mm in body length;
Enchiridium daidai sp. nov., photograph taken in life and eyespots observed in fixed state after being cleared in xylene. A
Enchiridium daidai sp. nov.,
Comparison of characters between five Enchiridium species in which marginal eyespots are distributed only anteriorly.
E. daidai sp. nov. | E. delicatum | E. gabriellae | E. magec | E. russoi | |
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Type locality | Off the coast of Bonomisaki, Kagoshima, Japan | East London, South Africa | São Sebastião Island, São Paulo, Brazil | North of El Balito, Tenerife, Canary Islands, Spain | Shelley Beach, East London, South Africa |
Dorsal coloration/ pattern: | |||||
Background color | Translucent | Light pale yellow | Transparent | Whitish to cream | Greyish yellow |
Spots or maculae on dorsal surface | None | None | None | Brown caramel spots, arranged more densely in the central region | Brown pigment spots spread especially in the central part |
Median line | None | Two yellow bands | None | A band composed of brown caramel spots | An ocher yellow band |
Fringed line | A thin orange line | None | None | None | None |
Reference | This study |
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We are very grateful to Mr Yuki Oya (Hokkaido University, Japan), Dr Naoto Jimi (National Institute of Polar Research, Japan), Dr Daisuke Uyeno (Kagoshima University, Japan), Ms Midori Matsuoka (Kagoshima University, Japan) and Mr Hiroki Kise (Ryukyu University, Japan) for help in collecting the specimens in Kagoshima and Okinawa. AT is thankful to Mr Yuki Oya for helpful comments about an earlier version of the manuscript. We thank Mr Daniel Whitmore for revision of the English language. This study was funded by Research Institute of Marine Invertebrates (FY 2019, No. 15 for AT).