Research Article |
Corresponding author: Takato Izumi ( iz.takato@gmail.com ) Academic editor: James Reimer
© 2018 Takato Izumi, Toshihiko Fujita.
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:
Izumi T, Fujita T (2018) Description of three new species of Scolanthus (Cnidaria, Anthozoa, Actiniaria, Edwardsiidae): first records of the genus from Japan. ZooKeys 794: 1-21. https://doi.org/10.3897/zookeys.794.25243
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Scolanthus is one genus of Edwardsiidae, a speciose family of burrowing worm-like sea anemones characterized by lacking a physa-like aboral end and by possessing nemathybomes on the whole body except at the distal end. This genus has been recorded worldwide, but there have been no specimens collected from Japan. In this study, we discovered four Scolanthus species in Japan for the first time: Scolanthus armatus (Carlgren, 1931) and Scolanthus kopepe sp. n. from the Ogasawara Islands, Scolanthus ena sp. n. from Ena Bay, Kanagawa, and Scolanthus isei sp. n. from Sugashima Island, Mie.
Aboral end, Ena Bay, nemathybome, Ogasawara Islands, sea anemone, edwardsiids, Sugashima Island, taxonomic key
The family Edwardsiidae is characterized by their worm-like bodies and is one of the major taxa in the order Actiniaria. Edwardsiidae contains approximately 90 species (
In Japan, 13 species of five genera of the family Edwardsiidae have been reported (Table
The genus Scolanthus was established by
The study of Edwardsiidae in Japanese waters has not progressed much, and additionally the genus Scolanthus has not been reported from Japan thus far. In this research, we report Scolanthus armatus (Carlgren, 1931) from the Ogasawara Islands, Japan, as well as discover and describe three new species from Japanese waters.
All Edwardsiidae sea anemones recorded in Japanese waters. Note some researchers recently have advocated that Metedwardsia akkeshi (and the genus Metedwardsia) do not belong to Edwardsiidae (
Species | Localities in Japan | Source |
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Edwardsia japonica Carlgren, 1931 | Misaki, Sagami Bay (Type locality) |
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Edwardsia octoradiata Carlgren, 1931 | Japan (Type locality) |
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Edwardsia arctica Carlgren, 1921 | 2300 m depth in Sea of Japan |
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Edwardsia sojabio Sanamyan & Sanamyan, 2013 | 500–3500 m in Sea of Japan (Type locality) |
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Paraedwardsia cretata (Stimpson, 1856) | Kagoshima Bay, Japan (Type locality) |
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Metedwardsia akkeshi (Uchida, 1932) | Akkeshi, Hokkaido (Type locality) |
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Tempuractis rinkai Izumi, Ise & Yanagi, 2018 | Misaki, Sado, Toba (Type localities) |
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Species below were only included in field guidebook | ||
Edwardsia sipunculoides (Stimpson, 1853) | Tohoku region |
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Edwardsianthus cf. pudica (Klunzinger, 1987) | Sagami Bay, Onagawa Bay (as E. japonica in Uchida, 1941 and 1965) |
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Edwardsianthus gilbertensis (Carlgren, 1931) | Kabira Bay, Okinawa |
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Edwardsia hantuensis England, 1987 | Boso Pninsula, Chiba |
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Edwardsia aff. norvegica (Carlgren, 1942) | Kii Peninsula, Wakayama |
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The specimens were dug out from sandy seafloors using a shovel and a sieve, or from rocky substrates using a chisel, by wading, snorkeling, or scuba diving. Sampling was carried out around Chichijima Island in the Ogasawara Islands, in Ena Bay in Kanagawa, and around Sugashima Island in Mie, Japan (Fig.
The examined specimens were deposited in the National Museum of Nature and Science, Tokyo (
Histological sections were made generally following standard protocols of
Cnidae were observed in the tentacle, actinopharynx, nemathybomes, column and filament; this genus does not have physa, so there are no cnidome data for the physa. Images of the cnidae were obtained by differential interference contrast microscopy (
Edwardsiidae with body divisible into scapus and capitulum. Proximal part of body rounded, provided with nemathybomes and never forming physa. Nemathybomes scattered or forming longitudinal lines on scapus. At least eight microcnemes. Tentacle typically 16 to 20 in adults, arranged hexamerously, octomerously or decamerously. Tentacles on inner cycle shorter than those on outer cycle. Retractor muscles relatively large, well developed, and diffused, restricted or circumscribed. Parietal muscles distinct, symmetrical, well developed. Cnidome; spirocysts, basitrichs, microbasic b- and p-mastigophores (revised from
Scolanthus callimorphus Gosse, 1853 by monotypy.
New Japanese name: Ashinashi-mushimodoki-ginchaku-zoku. This new Japanese name is constructed from “ashi-nashi” (which means no-foot) and “mushimodoki-ginchaku” (which means worm-like, the Japanese name for edwardsiids). “Ashi-nashi” is named after the most characteristic feature of this genus; no differentiated physa from scapus.
The genus Scolanthus currently contains seven valid species (
Edwardsia
armata
Carlgren, 1931: 2, figs 1–2;
Scolanthus
armatus
:
External anatomy. Column rough, ca. 40 mm in whole length, and ca. 5 mm in width, worm-like form, and the proximal part narrower to some extent. The column consists of capitulum, scapulus and scapus. The most proximal part of column capitulum, distinct, extremely short, whitish and semitransparent, but scapulus and scapus indistinct. The periderm-like cuticle, brownish orange with no pattern in color, covering the whole column except capitulum and tentacle but easily stripped off from epidermis (Fig.
External and internal views of Scolanthus armatus (
Internal anatomy. Eight perfect mesenteries, all macrocnemes. Four dorsal and ventral directives, and four lateral mesenteries not-paired with other macrocnemes, arranged in normal Edwardsia pattern (Fig.
Cnidome. Basitrichs (in all tissues), spirocysts (in tentacles), microbasic b-mastigophores (in filaments) (Table
Fiji (
New Japanese name: Ogasawara-ashinashi-mushimodoki. “Ogasawara” is the locality name. In Japanese waters, this species has been collected only in the Ogasawara Islands. “Ashinashi-mushimodoki” is short for the Japanese name of this genus (see the etymology of genus).
Scolanthus armatus was originally described by
In conclusion, we identified this specimen as S. armatus because of its similarity in almost all morphological features to the original description of the species. The slight difference observed in the cnidome may be simply individual variation.
Holotype.
External anatomy. Column rough, rugged and uneven, ca. 80 mm in whole length in holotype, and 10–15 mm in width, pipe-like in form both in living (Fig.
Internal anatomy. Eight perfect mesenteries, all macrocnemes. Four dorsal and ventral directives, and four lateral mesenteries not-paired with other macrocnemes (Figs
Cnidome. Spirocysts (in tentacles), basitrichs (in every tissue), microbasic b-mastigophores (in filament) and microbasic p-mastigophores (in actinopharynx, and filament) (Table
External and internal views of Scolanthus ena sp. n. (
Ena Bay, Kanagawa. Known only from the type locality.
The species epithet is named after the type locality, Ena Bay. The word “ena” is a noun in apposition. Origin of Japanese name: new Japanese name: taru-ashinashi-mushimodoki. “Taru” is the name of the collector of this new species.
Scolanthus ena sp. n. has 20 tentacles, as do Scolanthus ignotus (Carlgren, 1920) and S. isei sp. n.: other edwardsiid species have 16 tentacles (
This species is one of the biggest species in the genus Scolanthus: all previously reported species of nominal Scolanthus have bodies less than 80 mm in length (
Despite several sample collection surveys at Ena Bay, S. ena sp. n. was collected only once, and no specimens have been collected from any other locality. It is said by local people that the environment of Ena Bay has changed from several decades ago; the bottom of bay was previously rocky, and a muddy flat has formed in recent years by inflow of sediment. Considering some Scolanthus live in rocky habitats compared to other edwardsiids (e.g. S. isei sp. n.), the primary habitat of S. ena sp. n. might be rocky, and perhaps the numbers of individuals have decreased in Ena Bay by recent rapid changes in the environment. It is difficult, however, to examine this hypothesis because Edwardsiidae sea anemones living in or between rocks often cannot be collected easily even if there are many individuals present.
Even though there is only one specimen of S. ena, the character differences from other Scolanthus species make it obvious that this specimen is not a formerly described Scolanthus species. Examination of additional specimens in the future may help better delineate this species.
Holotype.
External anatomy. Column rough, rugged and uneven, ca. 30 mm in whole length in fixed holotype, and 10–12 mm in width, truncated cone-like form both in living and fixed (Fig.
Internal anatomy. Eight perfect mesenteries, all macrocnemes. Four dorsal and ventral directives, and four lateral mesenteries not paired with other macrocnemes (Fig.
Cnidome. Spirocysts (in tentacles), basitrichs (in all tissues), microbasic b-mastigophores (in actinopharynx) and microbasic p-mastigophores (in filament) (Table
External and internal views of Scolanthus isei sp. n. (
Southwest coast of Sugashima Island, Mie. Known only from the type locality.
The species name was named after Yuji Ise, the collector of both holotype and paratype specimens. Origin of Japanese name: New Japanese name: sugashima-gareba-ashinashi-mushimodoki; “gareba” means rocky seashore, the habitat where this species inhabits.
In terms of having 20 tentacles, Scolanthus isei sp. n. resembles S. ignotus and S. ena sp. n., while all other edwardsiid species have 16 tentacles (
Exceptionally for edwardsiids, this species inhabits in the cavities of the underside of boulders, adhering by their aboral end, on rocky seashores. Edwardsiids usually inhabit in sand or mud, and only two species had been reported from other different environments: Edwardsiella andrillae Daly, 2013 living on ice (
Holotype.
External anatomy. Column comparatively smooth, ca. 15–25 mm in whole length (25.0 mm in holotype), and 1–2 mm in width (1.8 mm in holotype), naked and extremely long and narrow pipe-like form both in living (Fig.
Internal anatomy. Eight perfect mesenteries, all macrocnemes. Four dorsal and ventral directives, and four lateral mesenteries not-paired with other macrocnemes, arranged in normal Edwardsia pattern (Figs
Cnidome. Spirocysts (in tentacles), basitrichs (in all tissues), microbasic b-mastigophores (in actinopharynx, column and filament) (Table
External and internal views of Scolanthus kopepe sp. n. A, B, F–I
Schematic representation of tentacular and mesenterial arrangements. A Scolanthus armatus and S. kopepe sp. n. B S. ena sp. n. C S. isei sp. n. Abbreviations: a, actinopharynx; bw: body wall; dd, dorsal directive; dlm, dorso-lateral mesentery; mi, microcneme; rm, retractor muscle; vd, ventral directive; vlm, ventro-lateral directive. White circles indicate the locus of tentacles.
Chichijima Island, Ogasawara. Known only from the type locality.
Most of the specimens were collected from Kopepe Coast. “Kopepe” is the name of the native people that used to live in the Ogasawara Islands, who had emigrated from the Gilbert Islands, now the Republic of Kiribati. The word “kopepe” is a noun in apposition. Origin of Japanese name: New Japanese name: kopepe-ashinashi-mushimodoki. “Kopepe” is originated from same coast.
This species resembles not only Scolanthus armatus but also S. scamiti Daly & Ljubenkov, 2008, S. triangulus Daly & Ljubenkov, 2008, S. curacaoensis (Pax, 1924), S. nidarosiensis (Carlgren, 1942) and S. callimorphus Gosse, 1853 in terms of having 16 tentacles (
Scolanthus kopepe sp. n. usually lives in coral sand, and prefers dark environments under large rocks.
Cnidae of Scolanthus species. A–E S. armatus (
Cnidae of four Japanese Scolanthus species. –absent. n = number of observed cnidae. A–R indicate figures of each kind of cnidae in Figure
Fig | Scolanthus armatus | Fig | S. ena sp. n. | Fig | S. isei sp. n. | Fig | S. kopepe sp. n. | |||||||||||||
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(specimen) |
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Length × Width (µm) | n | Length × Width (µm) | n | Length × Width (µm) | n | Length × Width (µm) | n | |||||||||||||
Tentacle | ||||||||||||||||||||
Spirocysts | A | 11.79–20.69 × 1.74–3.70 | 17.02 × 2.56 | 2.11 × 0.37 | 45 | F | 11.15–20.90 × 2.11–3.21 | 15.57 × 2.62 | 2.00 × 0.27 | 53 | J | 10.99–16.03 × 2.35–3.76 | 13.52 × 3.13 | 1.19 × 0.35 | 40 | O | 10.40–18.29 × 1.55–2.80 | 14.99 × 2.08 | 2.08 × 0.29 | 47 |
Basitrichs | 13.05–54.36 × 2.70–5.25 | 32.65 × 3.58 | 13.69 × 0.47 | 71 | 21.19–38.34 × 2.37–3.51 | 31.38 × 2.97 | 2.88 × 0.25 | 62 | small 10.05–12.97 × 1.82–2.77 | 11.58 × 2.33 | 0.70 × 0.31 | 22 | small 11.29–17.27 × 2.80–4.10 | 14.69 × 3.39 | 1.66 × 0.35 | 35 | ||||
large 19.10–26.60 × 3.04–4.52 | 22.54 × 3.71 | 1.64 × 0.33 | 59 | large 31.91–55.17 × 2.70–4.92 | 45.82 × 3.97 | 5.67 × 0.46 | 73 | |||||||||||||
Actinopharynx | ||||||||||||||||||||
Basitrichs | B | 19.44–39.57 × 2.67–4.75 | 29.10 × 3.57 | 4.30 × 0.51 | 40 | G | small 22.55–26.92 × 2.39–2.77 | 24.74 × 2.58 | 5.27 × 0.70 | 2 | K | 21.91–39.78 × 3.06–4.65 | 32.01 × 3.93 | 4.79 × 0.42 | 55 | P | small 14.94–28.33 × 2.32–3.77 | 20.27 × 3.13 | 1.80 × 0.35 | 22 |
large 37.46–51.19 × 2.95–4.62 | 43.98 × 3.90 | 3.12 × 0.32 | 65 | large 33.75–46.69 × 3.14–4.61 | 39.83 × 3.88 | 2.97 × 0.33 | 38 | |||||||||||||
Microbasic b–mastigophores | 22.17–34.53 × 3.48–5.19 | 28.32 × 4.09 | 2.81 × 0.40 | 42 | – | – | – | – | 25.14–38.09 × 4.24–6.43 | 31.02 × 5.45 | 2.85 × 0.41 | 49 | – | – | – | – | ||||
Microbasic p–mastigophores | – | – | – | – | 21.67–30.55 × 4.94–6.93 | 27.16 × 5.68 | 1.69 × 0.50 | 22 | – | – | – | – | – | – | – | – | ||||
Nemathybome | ||||||||||||||||||||
Basitrichs | C | 36.20–62.95 × 3.41–5.80 | 52.02 × 4.69 | 7.15 × 0.56 | 54 | H | small 9.74–15.15 × 2.20–3.05 | 12.07 × 2.53 | 1.38 × 0.23 | 16 | L | small 9.68–12.35 × 1.96–3.23 | 11.02 × 2.40 | 1.34 × 0.36 | 5 | Q | small 18.89–24.97 × 2.67–4.33 | 21.30 × 3.61 | 1.72 × 0.35 | 21 |
large 63.93–105.59 × 3.41–5.40 | 83.26 × 4.26 | 7.85 × 0.39 | 47 | large 62.84–84.98 × 3.76–6.16 | 73.91 × 4.87 | 9.76 × 0.60 | 52 | large 29.79–58.01 × 2.80–5.20 | 44.81 × 4.45 | 7.41 × 0.49 | 56 | |||||||||
Column | ||||||||||||||||||||
Basitrichs | D | 10.47–49.97 × 2.76–5.00 | 18.16 × 3.56 | 10.93 × 0.59 | 21 | – | – | – | – | – | – | – | – | – | R | 14.28–50.47 × 3.90–4.15 | 39.57 × 4.69 | 8.26 × 0.55 | 31 | |
Filament | ||||||||||||||||||||
Basitrichs | E | small 31.29–43.01 × 3.48–5.43 | 37.17 × 4.37 | 2.90 × 0.36 | 70 | I | 33.37–45.48 × 3.72–5.56 | 38.12 × 4.63 | 2.78 × 0.48 | 34 | M | small 18.18–26.59 × 2.28–3.61 | 21.68 × 3.02 | 2.32 × 0.36 | 21 | S | 19.57–29.72 × 2.70–4.25 | 23.32 × 3.60 | 2.11 × 0.31 | 66 |
large 95.62–114.46 × 2.67–4.16 | 102.12 × 3.64 | 8.28 × 0.69 | 3 | large 38.66–49.82 × 4.56–6.65 | 44.45 × 5.60 | 2.50 × 0.52 | 41 | |||||||||||||
Microbasic b–mastigophores | 33.80–45.69 × 5.21–7.90 | 43.19 × 6.54 | 2.14 × 0.71 | 21 | 34.26–39.98 × 5.11–6.55 | 36.85 × 5.80 | 1.91 × 0.46 | 12 | – | – | – | – | 32.30–39.62 × 5.24–7.77 | 35.88 × 6.22 | 1.68 × 0.51 | 57 | ||||
Microbasic p–mastigophores | – | – | – | – | 23.49–29.63 × 5.69–7.73 | 27.44 × 6.30 | 1.80 × 0.58 | 9 | 31.88–33.97 × 5.20–6.12 | 33.03 × 5.63 | 0.70 × 0.36 | 5 | – | – | – | – |
We are grateful to Masanori Taru (Toho University), who collected the Scolanthus ena sp. n. specimen, and Masaatsu Tanaka (Kagoshima University), who brought us the S. ena specimen. We thank Yuji Ise (Sugashima Marine Biological Laboratory, Nagoya University), and Sugashima Marine Biological Laboratory for helping in research of S. isei sp. n. We acknowledge Kensuke Yanagi (Coastal Branch of Natural History Museum and Institute of Chiba), who often guided our research. Finally, we thank James Davis Reimer (University of the Ryukyus) who revised and edited the English of this article.
This study was supported in part by a Sasakawa Scientific Research Grant from The Japan Science Society (No. 27-528), by JSPS KAKENHI Grant Number 17J03267, and by the Integrated Research Projects, “Biological Properties of Biodiversity Hot Spots in Japan” and “Geological, Biological, and Anthropological Histories in Relation to the Kuroshio Current”, conducted by the National Museum of Nature and Science.