Epitoniidae Berry, 1910 (1812)

Boreoscala greenlandica

(Perry, 1811)

Description:

Dimensions range from 15 mm - 61 mm high and 4 mm - 22 mm wide.
Width to height ratio: 0.27:1 ~ 0.36:1

Shell: attenuate, slender, elongate, conical; chalky, thick; small to medium. Exterior colour uniform greyish tan to chalky white, may be somewhat shiny or without lustre, costae white. About 16 whorls, moderately convex, joined; protoconch with 2-4 whorls smooth, usually eroded; teleoconch with 6-12 whorls moderately convex. Spire fairly attenuated. Suture moderately impressed, not deep, however Sars [1878:194] describes it deeply impressed. Imperforate. Approximately 9 - 34 costae on the body whorl. Costae are blade-like to ridge-like or strap like, thick, curved and may be expanded at the suture, sometimes nearly covering the intercostal spaces. Costae connected across the suture; slightly raised toward the suture; reflected. Intercostal spaces with flattened spiral striae. Interstices with approximately 8 - 10 spiral striae per whorl. Base with well developed basal ridge with flattened spiral striae below. Base with 3 - 8 spiral striae, Columella short and arched. Aperture subovate to circular, lip thickened; peristome thickened. Operculum black, horny, with a narrow brown border..

Distribution:

Probably cireumpolar [Clench & Turner, 1952]. Arctic Ocean near Point Barrow, Alaska, south to British Columbia, Canada; Siberia and northern Japan [Dushane, 1974], Hokkaido to Boso Peninsula, Japan; widely in the northern Pacific, Atlantic, Arctic Ocean [Nakayama, 2003]; Korean Peninsular [NIBR, 2019:327];Circumboreal, Scandinavia, Iceland [Brezinger et al., 2020]; Atlantic south to Long Island [Weil et al., 1999:12].

Habitat:

Shallow subtidal to upper continental slope [ Alf et al., 2020], subtidal to 650 m., sandy mud [Higo, et al., 1999].

Etymology:

.

Type Material:

Types. The whereabouts of Perry's types isunknown to us. We here restrict the type locality to Godthaab, southwest Greenland, one of the several localities listed by Posselt. The type specimen of Scalaria subulata Couthouy has been lost. The type of Scalaria planicosta Kiener is in the Paris Museum. A cotype of S. grénlandica var. crebricostata G. O. Sars is in the Oslo Museum, Oslo, Norway. for Boreoscala greenlandica (Perry, 1811). [Clench, W. J. & Turner, R. D., 1952b]
Types. The type of HE. greenlandicum lovenu A. Adams is probably in the British Museum. The type locality is here restricted to Vardo, Norway. [as Boreoscala greenlandica f. lovenii (A. Adams, 1856)]. [Clench, W. J. & Turner, R. D., 1952b]
Types. The holotype of Scalaria groenlandica ornata Friele and Grieg is in the Zoologisk Museum, Universitetet, Bergen, Norway, no. 21669. We figure the holotype from station 124 (N. Lat. 66°41'; EK. Long. 6°59') off Vest Fjorden, Norway. [as Boreoscala greenlandica norvegicum (Clench & R. D. Turner, 1952), Epitonium greenlandicum ornatum Friele and Grieg, Scalaria ornata Friele & Grieg, 1901]. [Clench, W. J. & Turner, R. D., 1952b]
Type: Scalaria greenlandicum Perry: Holotype lost (Clench & Turner, 1952: 320); type locality restricted to Godthaab, southwest Greenland (Clench & Turner, loc. cit.). Epitonium (Boreoscala) greenlandicum smithi MacNeil: Holotype USNM 499037; Inner Submarine Beach, Nome for Boreoscala greenlandica (Perry, 1811). [DuShane, H., 1979b]
Type not listed in source data for Boreoscala greenlandica (Perry, 1811) - Type Locality: Greenland (restricted by Clench and Turner, 1952: 320).. [Kazunori, H., 2009]
Types: S. greenlandica, not located; S. loveni, Cambridge Museum. for Boreoscala greenlandica (Perry, 1811) - Type Locality: S. greenlandica, restricted by Clench & Turner (1952:320) to Godthaab, SW Greenland.. [Bouchet, P. & Warén, A., 1986]
Type not indicated in text [as Cirsotrema (Boreoscala) greenlandicum (Perry, 1811)] - Type Locality: Greenland. [Nakayama, T., 2003]
Type not located by either Clench & Turner (1952: 320) or Bouchet & Warén (1986: 518). for Boreoscala greenlandica (Perry, 1811) - Type Locality: restricted to Godthaab, southwest Greenland, by Clench & Turner (loc. cit.).. [Brown, L. & Neville, B.D., 2015]
Type not located for Boreoscala greenlandica (Perry, 1811) - Type Locality: not designated.. [Brown, L. & Neville, B.D., 2015]
Cotype: ZMO figured by Clench & Turner (1952: 321, pl. 154, fig. 3). [as Scalaria grønlandica var. crebricostata Sars, 1878:] - Type Locality: Vadsø, Norway.. [Brown, L. & Neville, B.D., 2015]
Type: “ex museo nostro,” presumably the figured specimen. [as Turbo Clathrus groenlandicus Chemnitz, 1795 in Martini & Chemnitz, 1769-1829:] - Type Locality: Greenland.. [Brown, L. & Neville, B.D., 2015]
Type: Cambridge Museum (Bouchet & Warén 1986: 518) [as Scalaria lovenii A. Adams, 1856] - Type Locality: 'Ad litt. Scandinaviæ,' restricted to Vardö, Norway by Clench & Turner 1952: 322). [Brown, L. & Neville, B.D., 2015]
Type not located [as Scalaria grönlandica var. major Kobelt, 1888] - Type Locality: not designated. [Brown, L. & Neville, B.D., 2015]
Type not located [as Scalaria trevelyana var. major Locard 1897] - Type Locality: côtes océaniques de France. [Brown, L. & Neville, B.D., 2015]
Type not located [as Scalaria mediterranea var. major Locard in Locard & Caziot, 1899] - Type Locality: Corsica. [Brown, L. & Neville, B.D., 2015]
Type not located [as Opalia major Pallary, 1900] - Type Locality: Oran, Algeria. [Brown, L. & Neville, B.D., 2015]
Type: Zoologisk Museum, Universitetet, Bergen, Norway, no. 21669, figured by Clench & Turner (1952: 324). [as Scalaria goenlandica var. ornata Friele & Grieg, 1901:] - Type Locality: Norwegian North Atlantic Expedition 1876-78, S.S. Vöringen Station 124, off Trænen, Helgeland, Norway, 66o 41'N, 06o 59'E, 350 fms [= 640 m].. [Brown, L. & Neville, B.D., 2015]
Type: “Collect. du Mus.”; while Clench & Turner (1952: 320) report that the type is at the MNHN, it is not listed in the MNHN type catalog. [as Scalaria planicosta Kiener, 1838:] - Type Locality: not given.. [Brown, L. & Neville, B.D., 2015]
Type: “My own Collection. Cabinet of Bost. Soc. Nat. Hist. and of N. Y. Lyceum of Nat. Hist.” [ Couthouy, 1838] [as Scalaria subulata Couthouy, 1838:] - Type Locality: “Mass. Bay, vicinity of Cape Ann. [Brown, L. & Neville, B.D., 2015]
Syntype: KPM-NG0102035 [as Borcoscala groenlandium japonicum (SHIKAMA, 1962)] - Type Locality: East to southeast of Choshi (eastern central Honshü). [Higo, S., Callomon, P. & Goto, Y., 2001]

SpeciesImage

Dall, W. H. (1925). Plate 22. Fig. 2.

Boreoscala greenlandica Perry, alt. 52 mm

Distribution

as listed in source literature

Sowerby, J. & Sowerby, J. de C. (1823).

Shown in text as Boreoscala subulata (Couthouy, 1838)

Species Image

Plate cccxc. Fig. 1.

Species Image

Plate cccxc. Fig. 1.

Species Image

pp: 125.

Couthouy, J. P. (1838).

Shown in text as Scalaria subulata Couthouy, 1838

Species Image

Plate 3. Fig. 4.

Species Image

pp: 93-94.

Species Image

pp: 95-96.

Kiener, L. C., & Fischer, P. (1838b).

Shown in text as Scalaria planicosta Kiener, 1838

Species Image

Plate VII. Fig. 21.

Species Image

Plate VII. Fig. 21.

Species Image

pp: 18.

16. Flat-ribbed Wentletrap-Scalaria planicosta Nobis
(Museum collection)
(Pl. VII, fig. 21)
Shell elongate, turreted, reddish or greyish, apex acute; whorls somewhat convex, contiguous, with flattened, distant longitudinal ribs; interspaces transversely striated.
Shell elongate, turreted, not umbilicate, very pointed at the apex, composed of 10- slightly convex, contiguous whorls, bearing 13-14 rather thick longitudinal ribs, generally flattened and widely spaced, several of which occasionally develop into varices. These ribs join superficially at the deep suture and form undulating bands covering the entire shell. Regularly spaced transverse striae, equal amongst themselves and much less prominent than the ribs, are visible in the interspaces. Aperture subrounded, with thickened, slightly reflected margins. Colour a pleasing reddish or pale grey background against which the ribs, all white, stand out distinctly.
Length 22 lines.
Habitat. Not stated.
This attractive shell, belonging to the Museum collection, is distinguished by its ribs, which are flatter and more numerous than those of Scalaria communis Scalaria communis. It also differs from that species in the regular transverse grooves described above.

Sowerby, G. B. II. (1844).

Shown in text as Scalaria greenlandica G. Perry, 1811

Species Image

Plate xxxiv. Fig. 110.

Species Image

pp: 101.

Shell elongate, thick, spiral grooved; with contiguous whorls, barely prominent; varices reclining; unequal, expanding toward the suture, bent back, and elevated toward the nearest whorl; aperture somewhat triangular, with a grayish-dark colour between the varices.

Species Image

Plate xxxiv. Fig. 105.

Reeve, L.A. (1873).

Shown in text as Scalaria greenlandica G. Perry, 1811

Species Image

Plate xiv. Fig. 107a.

Species Image

Plate xiv. Fig. 107b.

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Sars, G. O. (1878).

Shown in text as Scalaria greenlandica var. crebricostata G. O. Sars, 1878

Species Image

Plate 23. Fig. 1.

Species Image

pp: 194.

c. var. crebricostata (Pl. 23, fig. 1).
Shell narrower, spire more exserted and gradually attenuated; whorls fairly convex; suture deeply impressed; longitudinal folds very narrow, about 15 on the last whorl; spiral lines very distinct. Length 26 mm.

Sars, G. O. (1878).

Shown in text as Scalaria greenlandica G. Perry, 1811

Species Image

Plate 10. Fig. 15a-b.

Species Image

pp: 194.

99. Scalaria groenlandica, Chemn.
a. typical form (Pl. 10, figs. 15a—n¢?æ$—n¢?G=cÙƒ

Sars, G. O. (1878).

Shown in text as Scalaria greenlandica var. loveni A. Adams, 1856

Species Image

Plate 10. Fig. 16.

Species Image

b. var. Lovéni (Pl. 10, fig. 16).
Shell generally larger and more swollen; folds of the last 4

Tryon, G.W. (1887).

Shown in text as Scalaria groenlandica Chemnitz, 1795 in Martini & Chemnitz

Species Image

Plate 16. Fig. 91.

Species Image

pp: 76.

Species Image

pp: 76.

Dall, W. H. (1889b).

Species Image

Plate LXXII. Fig. 10.

Species Image

pp: 124.

Clessin, S. (1897).

Species Image

pp: 65-66.

Shell solid, white, opaque, somewhat shiny, narrowly turreted, with a rather attenuated spire; whorls 9

Species Image

Plate 17. Fig. 1.

Species Image

Plate 17. Fig. 3.

Dall, W. H. (1925).

Shown in text as Boreoscala greenlandica Perry, 1811

Species Image

Plate 22. Fig. 2.

Boreoscala greenlandica Perry, alt. 52 mm

Species Image

pp: 6.

Clench, W. J. & Turner, R. D. (1952b).

Shown in text as Boreoscala greenlandica norvegicum (Clench & R. D. Turner, 1952), Epitonium greenlandicum ornatum Friele and Grieg, Scalaria ornata Friele & Grieg, 1901

Species Image

pp: 323. Plate 155. Fig. 1.

Plate 155
Fig. 1. Epitonium greenlandicum ornatum Friele and Grieg
(= E. norvegicum Clench and Turner) from off Vest Fjorden, Norway. Holotype (2.8x).

Clench, W. J. & Turner, R. D. (1952b).

Shown in text as Boreoscala greenlandica f. lovenii (A. Adams, 1856)

Species Image

pp: 323. Plate 155. Fig. 2.

Epitonium greenlandicum loventt A. Adams from Browns Bank off Cape Sable, Nova Scotia in 40 fathoms (2.8x).

Clench, W. J. & Turner, R. D. (1952b).

Species Image

pp: 321. Plate 154. Fig. 1-3.

Epitonium greenlandicum Perry
Fig. 1. From Middle Bank, off Massachusetts in 35 fathoms (3x).
Fig. 2. From Swampscott, Massachusetts (2.8x).
Fig. 3. Epitonium greenlandicum crebricostatum Sars (= greenlandicum Perry), from Vadsé, Norway. Cotype (3.5x)

DuShane, H. (1979b).

Shown in text as Scalaria polita G.B. Sowerby, II, 1844: ~ Epitonium appressicostatum Dall, 1917; ~ Epitonium (Nitidiscala) pedroanum Willett, 1932, ~ Epitonium implicatum Dall & Ochsner, 1928

Species Image

pp: 103. Fig. 21.

Scalaria polita Sowerby, 1844 (after Sowerby, 1844). Holotype presumably lost; not found at BM(NH) by M. Keen in 1965. No dimensions given

Species Image

pp: 103. Fig. 22.

Epitonium (Nitidiscala) pedroanum Willett, 1932. Holotype, LACM 1036; length, 1 1.7 mm; width, 3.7 mm X 4.7

Species Image

pp: 103. Fig. 23-24.

Figure 23: Epitonium implicatum Dall & Ochsner, 1928. Holo- type, CAS(GTC) 2932; length, 18 mm; width, 6 mm X 3.5

Figure 24: Epitonium appressicostatum Dall, 191 7. Syntype, US NM 59334; length, 14 mm; width, 4.5 mm X4.5

DuShane, H. (1979b).

Species Image

pp: 103. Fig. 18.

Figure 18: Boreoscala greenlandica (Perry, 181 1). Greenland From Perry, 181 1 :
pit. 28, fig. 8

Species Image

pp: 103. Fig. 19-20.

Fig 19: Boreoscala greenlandica (Perry, 1811). Queen Char- lotte Islands, B. C, Canada, from 146 m, rock and sand sub- strate, collected by Bernard and Quayle, October, 1966; length, 39 mm; width, 14 mm (DuShane Collection) X 3

Figure 20: Boreoscala greenlandica smithi (MacNeill, 1943). Hol- otype, USNM 499037; length, 14.6 mm; width, 8 mm; Pliocene X 3

Bouchet, P. & Warén, A. (1986).

Species Image

pp: 518. Fig. 1210.

Figs 1210-1213. Epitonium greenlandicum. 1210, Greenland, 93 mm. 121, 66°23N, 20°15W, 80 m, 20.0 mm. 1212, very young specimen, Trondheimfjord, Norway, 1.95 mm. 1213, recently hatched juvenile. Bergen area. Norway. 1.4 mm.

Species Image

pp: 518.

Higo, S., Callomon, P. & Goto, Y. (2001).

Shown in text as Borcoscala groenlandium japonicum (SHIKAMA, 1962)

Species Image

pp: 54. Fig. G1877s.

G1877s
23.7
Borcoscala groenlandium
japonicum (SHIKAMA, 1962)
ST: KPM-NG0102035

Nakayama, T. (2003).

Shown in text as Cirsotrema (Boreoscala) greenlandicum (Perry, 1811)

Species Image

pp: 111. Plate 4. Fig. 20-22.

Cirsotrema (Boreoscala) greenlandicum (Perry, 1811)
height 60.0 mm; breadth 22.0 mm (NA); fig. 22, base

Kazunori, H. (2009).

Species Image

pp: 274. Fig. 138.

138. Boreoscala greenlandica, WA05-EF250D (empty shell).

Species Image

pp: 275.

Synonymy:

Borcoscala groenlandium japonicum (SHIKAMA, 1962) [in Higo, S., Callomon, P. & Goto, Y., 2001].
Boreoscala greenlandica f. lovenii (A. Adams, 1856) [in Clench, W. J. & Turner, R. D., 1952b].
Boreoscala greenlandica norvegicum (Clench & R. D. Turner, 1952), Epitonium greenlandicum ornatum Friele and Grieg, Scalaria ornata Friele & Grieg, 1901 [in Clench, W. J. & Turner, R. D., 1952b].
Boreoscala greenlandica Perry, 1811 [in Dall, W. H., 1925].
Boreoscala groenlandica (Chemnitz, 1795) [in Gul'bin, V.V., 2004].
Boreoscala subulata (Couthouy, 1838) [in Sowerby, J. & Sowerby, J. de C., 1823].
Cirsotrema (Boreoscala) greenlandicum (Perry, 1811) [in Nakayama, T., 2003].
Epitonium greenlandicum norvegicum Clench & Turner, 1952: [in Brown, L. & Neville, B.D., 2015].
Opalia major Pallary, 1900 [in Brown, L. & Neville, B.D., 2015].
Scalaria goenlandica var. ornata Friele & Grieg, 1901: [in Brown, L. & Neville, B.D., 2015].
Scalaria greenlandica G. Perry, 1811 [in Reeve, L. A., 1873].
Scalaria greenlandica G. Perry, 1811 [in Sars, G. O., 1878].
Scalaria greenlandica G. Perry, 1811 [in Sowerby, G. B. II., 1844].
Scalaria greenlandica var. crebricostata G. O. Sars, 1878 [in Sars, G. O., 1878].
Scalaria greenlandica var. loveni A. Adams, 1856 [in Sars, G. O., 1878].
Scalaria groenlandica Chemnitz, 1795 in Martini & Chemnitz [in Tryon, G.W., 1887].
Scalaria grønlandica var. crebricostata Sars, 1878: [in Brown, L. & Neville, B.D., 2015].
Scalaria grönlandica var. major Kobelt, 1888 [in Brown, L. & Neville, B.D., 2015].
Scalaria lovenii A. Adams, 1856 [in Brown, L. & Neville, B.D., 2015].
Scalaria lovenii A. Adams, 1856 [in Tryon, G.W., 1887].
Scalaria mediterranea var. major Locard in Locard & Caziot, 1899 [in Brown, L. & Neville, B.D., 2015].
Scalaria planicosta Kiener, 1838 [in Kiener, L. C. & Fischer, P., 1838b].
Scalaria planicosta Kiener, 1838: [in Brown, L. & Neville, B.D., 2015].
Scalaria subulata Couthouy, 1838 [in Couthouy, J. P., 1838].
Scalaria subulata Couthouy, 1838: [in Brown, L. & Neville, B.D., 2015].
Scalaria trevelyana var. major Locard 1897 [in Brown, L. & Neville, B.D., 2015].
Turbo Clathrus groenlandicus Chemnitz, 1795 in Martini & Chemnitz, 1769-1829: [in Brown, L. & Neville, B.D., 2015].

Source Literature:

Alf, A., Brenzinger, B., Haszprunar, G., Schrödl, M. & Schwabe, E. (2020). A Guide to Marine Molluscs of Europe. Conch Books. Conch Books
Bouchet, P. & Warén, A. (1986). Revision of the Northeast Atlantic bathyal and abyssal Aclididae Eulimidae, Epitonidae (Mollusca, Gastropoda).. Bollettino Malacologico.. suppl. 2. https://www.biodiversitylibrary.org/bibliography/140762 [Accessed 21 September 2023]
Brown, L. & Neville, B.D. (2015). Catalog of the recent taxa of the families Epitoniidae and Nystiellidae (Mollusca: Gastropoda) with a bibliography of the descriptive and systematic literature. Zootaxa. 3907(1). Magnolia Press Auckland, New Zealand. http://dx.doi.org/10.11646/zootaxa.3907.1.1 [Accessed 22 October 2023]
Clench, W. J. & Turner, R. D. (1952b). The Genus Epitonium in the Western Atlantic pt 2. Jonsonia. 2(31). The Department of Mollusks Museum of Comparative Zoology Harvard University. https://www.biodiversitylibrary.org/page/41155986#page/303/mode/1up [Accessed 25 June 2023]
Clessin, S. (1897). Die Familie der Scalariidae. In W. Kobelt (Ed.), Systematisches Conchylien-Cabinet von Martini und Chemnitz. Neu herausgegeben und vervollständigt. 2(13). Bauer & Raspe, NürnbergBauer & Raspe, Nürnberg. https://www.biodiversitylibrary.org/page/34259905 [Accessed 21 June 2025]
Couthouy, J. P. (1838). Descriptions of new species of Mollusca and shells, and remarks on several Polypii, found in Massachussets Bay.. Boston Journal of Natural History. 2(1). https://biodiversitylibrary.org/page/32266711 [Accessed 4 January 2026]
Dall, W. H. (1889b). Report on the Mollusca. Part II. Gastropoda and Scaphopoda. In Reports on the results of dredging by the U.S. Coast Survey Steamer "Blake". United States National Museum Bulletin. 37. Government Printing Office. https://ia800603.us.archive.org/22/items/bulletinunit71889unit/bulletinunit71889unit.pdf?utm_source=chatgpt.com [Accessed 2 May 2026]
Dall, W. H. (1925). Illustrations of unfigured types of shells in the collections of the United States National Museum. Proceedings of the United States National Museum. 66(2554). https://www.biodiversitylibrary.org/page/7738736 [Accessed 18 September 2025]
Durham, J. W. (1937). Gastropods of the Family Epitoniidae from Mesozoic and Cenozoic Rocks of the West Coast of North America, including One New Species by F. E. Turner and One by R. A. Bramkamp.. Journal of Paleontology. 11(6). http://www.jstor.org/stable/1298354 [Accessed 9 May 2026]
DuShane, H. (1979b). The Family Epitoniidae in the Northeastern Pacific. The Veliger. 22(2). California Malacozoological Society, Inc.. https://www.biodiversitylibrary.org/item/134937#page/7/mode/1up [Accessed 28 June 2023]
Gulbin, V. V. (2010). Review of the Shell-bearing Gastropods in the Russian Waters of the East Sea. II. Caenogastropoda: Sorbeoconcha, Hypsogastropoda. Korean Journal of Malacology. 26(2). https://koreascience.kr/article/JAKO201018860403872.pdf [Accessed 20 September 2025]
Higo, S., Callomon, P. & Goto, Y. (1999). Catalogue and Bibliography of the Marine Shell-Bearing Mollusca of Japan.. https://www.researchgate.net/publication/266375656_Catalogue_and_Bibliography_of_the_Marine_Shell-Bearing_Mollusca_of_Japan [Accessed 25 July 2025]
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Kazunori, H. (2009). Upper bathyal gastropods of of the Pacific coast of northern Honshu, Japan, chiefly collected by R/V Wakataka-Maru. National Museum of Nature and Science Monographs. 29. https://www.researchgate.net/publication/285740752 [Accessed 25 August 2023]
Kiener, L. C. & Fischer, P. (1838b). Genre Scalaire. Spécies général et iconographie des coquilles vivantes comprenant la collection du Muséum d’histoire naturelle de Paris : la collection Lamarck, celle du prince Masséna (appartenant maintenant a M.B. Delessert) et les découvertes récentes des voyageurs. 10. J.B.Baillière et Fils. http://www.biodiversitylibrary.org/item/19879 [Accessed 11 May 2026]
Nakayama, T. (2003). A Review of Northwest Pacific Epitoniids. Monographs of Marine Mollusca. 6. Backhuys Publishers
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Sars, G. O. (1878). Mollusca regionis arcticae Norvegiae. In Bidrag til kundskaben om Norges arktiske fauna. I: Oversigt over de i Norges arktiske region forekommende bløddyr. Brøgger
Sowerby, G. B. II. (1844). Monograph of the genus Scalaria.. Thesaurus conchyliorum, or monographs of genera of shells.. 1 (4). London, privately published.. http://biodiversitylibrary.org/page/11076419 [Accessed 22 June 2023]
Sowerby, J. & Sowerby, J. de C. (1823). The mineral conchology of Great Britain Vol IV. The mineral conchology of Great Britain; or, Coloured figures and descriptions of those remains of testaceous animals or shells, which have been preserved at various times and depths in the earth. Vol. IV.. IV. Privately published by author, London,. https://www.biodiversitylibrary.org/page/14392771 [Accessed 24 August 2023]
Tryon, G.W. (1887). Manual of conchology, structural and systematic with illustrations of the species. IX. Academy of Natural Sciences. https://www.biodiversitylibrary.org/page/11026889 [Accessed 20 September 2025]
Weil, A., Brown, L. & Neville, B. (1999). The Wentletrap Book. Evolver
WoRMS Editorial Board, (2025). World Register of Marine Species. https://www.marinespecies.org [Accessed 8 November 2025]

WoRMS direct page link: Open WoRMS record