Evolution of Janthina and Recluzia (Mollusca: Gastropoda: Epitoniidae) Author Beu, Alan G. text Records of the Australian Museum 2017 Rec. Aust. Mus. 2017-08-23 69 3 119 222 http://dx.doi.org/10.3853/j.2201-4349.69.2017.1666 journal article 228171 10.3853/j.2201-4349.69.2017.1666 d63960a1-e0c5-4b97-8e7d-f8be80d376d6 2201-4349 4677010 08B086EB-8D24-4FD0-975A-E045E2596BF1 Recluzia lutea ( Bennett, 1840 ) Figs 2S–T , 36E–H, K–P , 37 Janthina lutea Bennett, 1840: 298 (refers to description by Bennett, 1840: 63 ). Janthina turrita “von dem Busch ” Philippi, 1849: 15 ; H. Adams & A.Adams, 1854: 87; Mörch, 1860: 284 ; Tryon, 1887: 39 . Recluzia jehennei Petit de la Saussaye, 1853: 118 , pl. 5, fig. 3; H. Adams & A. Adams, 1854: 87; Mörch, 1860: 283 ; Küster, 1868: 11 , pl. 1, fig. 15; Tryon, 1887: 39 , pl. 10, fig. 28; Smith, 1910: 202 ; Fisher-Piette, 1950: 14; Bosch et al ., 1995: 111 , fig. 443. Recluzia rollandiana Petit de la Saussaye, 1853: 119 , pl. 5, fig. 2; H. Adams & A. Adams, 1854: 87, pl. 69, fig. 2; Chenu, 1859: 119 , fig. 520; Mörch, 1860: 284 ; Küster, 1868: 12 , pl. 1, fig. 14; Tryon, 1887: 38 , pl. 10, figs 25–27; Abbott, 1963: 151 ; Habe, 1964: 48 , pl. 14, fig. 7; Habe & Kosuge, 1966: 102, pl. 40, fig. 10; Bennett, 1966: 43 , pl. 11, fig. 1; Abbott, 1968: 92 , fig. 4; Cernohorsky, 1972: 198 , pl. 56, fig. 11; Abbott, 1974: 113 , fig. 1182; Powell, 1976: 107 ; Powell, 1979: 254 , pl. 48, fig. 23; Colman, 1986: 3 , text-fig.; Rios, 1994: 102 , pl. 33, fig. 417; Spencer & Willan, 1996: 24; Smith, 1998: 813 , fig. 15.151B; Spencer et al ., 2009: 206 ; Raven & Bracegirdle, 2010: 28, text-fig. Ianthina lutea Bennett. – Benson, 1860: 413 . Recluzia bensoni A. Adams, 1861: 402 ; Yen, 1942: 219 , pl. 19, fig. 122; Tryon, 1887: 39 . Recluzia montrouzieri Souverbie, 1871: 334 ; Souverbie, 1872: 20 : 57 , pl. 1, fig. 8; Habe & Tokioka, 1953: 57, fig. 12. Lymnaea ? palmeri Dall, 1871: 135 . Recluzia globosa E.A. Smith, 1876: 551 , pl. 30, fig. 8; Tryon, 1887: 39 , pl. 10, fig. 29. Recluzia rollandiana var. β annamitica Wattebled, 1886: 67 , pl. 3, figs 4a–b; Tryon, 1887: 38 , pl. 10, fig. 26. Recluzia lutea (Bennett) .– Oliver, 1915: 525 ; Hedley, 1918 : M62; Powell, 1924: 285 ; Powell, 1937: 74 ; Powell, 1946: 77 ; Powell, 1957: 98 ; Powell, 1962: 91 ; Rehder, 1980: 53 , pl. 7, fig. 6; Boone, 1984: 8 ; Higo et al ., 1999: 174 ; Okutani, 2000: 319 , pl. 158, fig. 5; Poppe, 2008: 722 , pl. 306, fig. 2. Recluzia palmeri (Dall) .– Dall, 1925: 25 , pl. 17, fig. 8; Keen, 1971: 443 , fig. 690; Abbott, 1974: 114 ; Abbott & Dance, 1982: 70, bottom right fig.; Poorman, 1980: 183 . Recluzia effusa “Martens ” Thiele, 1928: 78 , footnote, fig. 2 (shell not illustrated). Recluzia hargravesi Cox. – Allan, 1950: 95 , text-fig. 22.6; Wilson, 1993: 281 , pl. 44, figs 35A–B (misidentification as R. johnii Holten, 1802 ). Recluzia rollandiana bensoni A.Adams. – Kuroda et al ., 1971: 245 , pl. 62, fig. 16. Recluzia cf. jehennei Petit. – Churchill et al ., 2011a: 802 , fig. 1 (upper); Churchill et al. , 2011b: 441 , figs 1A–F. Type material . No type material of Janthina lutea is present in any museums the writer has consulted, and there is nothing in Bennett’s (1840) work to indicate that he retained any specimens. Bennett’s (1840: 63) description stated merely that this is an elongate, yellow species of Janthina with narrower whorls than J. janthina ; a neotype therefore is required to associate this name unambiguously with the present Recluzia species. The type locality is 2°53'S 174°55'E ( Bennett, 1840: 62 ), a short distance southwest of the Phoenix Islands , Kiribati , central western Pacific Ocean. However, as Recluzia lutea is pelagic and almost cosmopolitan, occurring in all world tropical and temperate seas, selection of a neotype from near the original type locality seems unimportant in this case, particularly as almost no material from the central Pacific has been observed by the writer. The writer also has been unable to trace any type material of Janthina turrita . Little that is diagnostic is available for this species, but the name “turrita” suggests that it applies to a Recluzia species rather than to Janthina , and the dimensions (H 14.5, D 10 mm ) rule out a position in Janthina . The name was not illustrated and has been referred to again only by Tryon (1887: 39) , who listed it among inadequately known species. The original description stated that the shell is fusiform, turreted, dark, smooth, with well-rounded whorls, deep sutures, and a short spire. The outer lip is straight and recedes strongly. The upper whorl surface is blue, and the base is pale reddish shading to rust red. The locality is unknown. Apart from the mysteries of the receding (incomplete?) lip and the colour, there seems to be little else this could refer to other than a Recluzia species. The late Dr Rudolf Kilias informed the writer that Dr Gerhard von dem Busch was a physician in Bremen . Enquiry at Übersee Museum Bremen revealed that von dem Busch’s collection is indeed located there. Janthina turrita is listed in von dem Busch’s manuscript catalogue with the locality “Adelaide”. However, the specimen is not present (T. Kruckow, Übersee Museum Bremen pers. comm. 05 Mar 1975 ), presumably because it was sent to Philippi. The writer has also been informed by N. Bahamonde (Museo Nacional de Historia Natural, Santiago , pers. comm. 25 Mar 1975 ) that no material identified as Janthina turrita is present there in Philippi’s collection. C. Zorn ( ZMB pers. comm. 30 Jan 2016 ) has also informed the writer that no material bearing this name is present in ZMB. Therefore, this name also requires a neotype . Recluzia jehennei , one figured syntype MNHN25667 ( Figs 36F, G ), listed by Fisher-Piette (1950: 14) as the holotype ; “this shell was found in the open ocean, in the Gulf of Arabia, by Commandant Jehenne” (translation from Petit de la Saussaye, 1853: 119 ). Recluzia rollandiana , no type material present in MNHN. The description by Petit de la Saussaye (1853: 120) indicated that the specimen remained “belonging to M Rolland de Roquan”, and so presumably is long lost; from “environs de Mazatlan” (“later identified as actually from New Caledonia ”; Keen 1971: 903 ). The remaining syntype of Recluzia jehennei , MNHN25667 ( Figs 36F, G ) is here designated the neotype of Janthina lutea Bennett, 1840 , the neotype of Janthina turrita Philippi, 1849 and the neotype of Recluzia rollandiana Petit de la Saussaye, 1853 , to refer these names unambiguously to the present species rather than to R. johnii . Recluzia bensoni , holotype NHMUK1878.1.28.394, labelled “ China Sea, R. Benson”, a small, narrow, juvenile specimen of Recluzia lutea , illustrated by Yen (1942: 219 , pl. 20, fig. 122). Yen’s illustration suggests that R. bensoni is possibly part of the variation of R. johnii , but the dimensions plot at the base of the field of R. lutea in Fig. 34 . Recluzia montrouzieri , one specimen in MNHN presented by the author was formerly considered to be a syntype but, as with Janthina capreolata , the type material was stated by Souverbie (1871) to be in MHNB, and the MNHN specimen is not a type; two syntypes MHNB2004.TY.163, from Art Island, New Caledonia ; photograph of syntypes sent by Laurent Charles, MHNB and Virginie Héros, MNHN (pers. comm. 21 Sep 2012 ). Lymnaea palmeri , holotype USNM56411 ( Dall, 1925: 25 , pl. 17, fig. 8), from the delta of the Taqui River, near Guaymas, Mexico , head of the Gulf of California. Abbott & Dance (1982: 70, bottom right fig.) illustrated the holotype of Lymnaea palmeri clearly. This specimen agrees in shape with that of the taller specimens of Recluzia lutea , but its original stated dimensions placed it within the field of R. johnii in Fig. 34 . The photographs ( Figs 36E, H ) sent by E. Strong (USNM pers. comm. 20 Sep 2012 ) show that the stated dimensions ( Keen, 1971: 443 ) were probably inaccurate, but they have been retained in Table 13 and Fig. 35 . The overall appearance is accepted here as confirming that R. palmeri is a synonym of R. lutea . Recluzia globosa , holotype NHMUK1876.1.10.119, labelled “Tarawa, Gilbert Group, received from John Brazier Esq. per Mr Henry Adams”, another small, very short specimen. All small specimens of R. lutea are very short compared with adults, and the dimensions of the holotype fall within the range of variation of R. lutea ( Fig. 35 ). Recluzia rollandiana var. annamitica , two syntypes MNHN25668 (one in Figs 36K–L ), from beach at Thuan-an, near Hué, Vietnam . Recluzia effusa holotype ZMB/Moll-13704 (E. Strong, USNM, pers. comm. 20 Sep 2012 ), not seen; from “chinesischen See”. The status of this name is not clear, as the shell is poorly preserved, but it is assumed here to be a further synonym of the more common Recluzia species. Other material examined . Present-day specimens from Australia and New Zealand: Australia : New South Wales : Dingo Beach, Cape Gloucester ( AMS ); Catherine Hill Bay ( AMS C11333); Port Macquarie ( NMV ); Collaroy Beach, Sydney ( AMS ); Cronulla, S of Sydney ( AMS C75719). Queensland : Michaelmas Cay, off Cairns ( AMS C53537); King’s Beach, Caloundra ( NMV , AMS ); Lady Elliott I. ( AMS C69170). Tasmania : Black Rocks, Richmond River ( AMS C76081). Kermadec Islands: Raoul I. ( SAMA ; AMS C36656; GNS WM5654, 1; NMNZ M202865,5; M202866, 1; M200986, 3; M202867, 1; M214384,20). New Zealand : Twilight Beach , Cape Reinga ( NMNZ M100807 , 2 ); N end of Ninety Mile Beach ( NMNZ M117600 , 6 ; M117826, 7); S of Scott Pt. , Ninety Mile Beach ( NMNZ M117256 , 20 ); S of the Bluff , Ninety Mile Beach ( NMNZ M277745 , 3 ); Great Exhibition Bay , Mar 1998 ( GNS RM6748 , 4 ; NMNZ M155930 , 59 ; P. Poortman colln., 20); Rarawa , Great Exhibition Bay ( NMNZ M081519 , 1 ); Taupiri Bay , N of Whangarei ( NMNZ M081518 , 26 ); Laings Beach , Mangawai , Hauraki Gulf ( AWM ); Mangawai Heads ( NMNZ M277744 , 1 ); Tryphena , Great Barrier Island ( Powell 1924: 285 ; in AWM ); Whangamata , Bay of Plenty ( AWM ). Figure 37. Recluzia lutea (Bennett) , NMNZ M081518, small specimens and larval shells; beach, Taupiri Bay, Northland, New Zealand; SEM. (A–B) apical views of shell and protoconch. (C–D , G) apertural view of incomplete specimen with 10 larval shells attached to dried float material; C, lateral view of protoconch of large specimen; D, whole shell; G, larval shell attached by dried float material to large specimen below its protoconch. (E–F) third specimen, lateral views of shell and protoconch. Scale bars: A, D, E, 1 mm; B, F, 200 µm; C, G, 300 µm. Distribution . Recluzia lutea occurs uncommonly throughout the world tropical and temperate ocean, but its limits are very poorly known. Large rafts of specimens are cast ashore in some locations, but much less frequently than for all Janthina species. In New Zealand , R. lutea is recorded only from the northeastern North Island warm-water region, in eastern Northland and the Bay of Plenty , as far southeast as Waihau Bay, easternmost Bay of Plenty . Much the largest lots have been seen from Great Exhibition Bay and Ninety Mile Beach in the northernmost North Island, where specimens are sometimes cast ashore in large numbers after unusually persistent onshore winds. Thompson (1991) gave a popular account of wash-ups of “about three hundred” specimens during Easter 1991 at Cape Maria van Diemen and Great Exhibition Bay, northernmost North Island. Large numbers of specimens also are cast shore from time to time all around eastern, northern and Western Australia. Specimens are also recorded from throughout the Indo-West Pacific province as far north as central Japan , in the Red Sea, and rarely in the eastern Pacific and the Atlantic Ocean. Dimensions . See Table 13 . Diagnosis . Moderate-sized (H to 36, D to 22 mm ), larger than Janthina exigua and J. umbilicata but smaller than all other Janthina species and Recluzia johnii ; taller and narrower than all Janthina species, but shorter and wider than R. johnii ; mean height: diameter 1.44, mean diameter: spire height 1.51. Teleoconch of 4.5–5.5 whorls; whorls strongly and evenly inflated, of almost circular cross-section, wider than in R. johnii ; suture deeply impressed; without obvious sculpture. Aperture slightly flared anteriorly in many large specimens. Lays smooth, narrow, cylindrical to weakly clubshaped, yellow egg capsules on underside of float. Remarks . Recluzia lutea is superficially smooth and lightly polished when the periostracum is removed, but most specimens are found with the thin, pale brown, matt periostracum still at least partly adhering. Recluzia lutea has strongly and evenly inflated whorls and a much more deeply impressed suture than in any Janthina species. Weak axial and spiral grooves and ridges occur on many specimens, varying individually. The outer lip is vertical and straight, i.e., with no sinus in the lip, and the inner lip is narrowly but variably reflected over the narrow umbilicus. Bennett’s (1840) description of Janthina lutea is quite adequate to make this name available, although which species he had is not determinable from the description. Bennett (1840: 298) stated “In the course of the narrative I have had occasion to mention a yellow and very rare species of this genus [ Janthina ]: and for which I would propose the name Janthina lutea ”. This is a reference to his earlier description ( Bennett, 1840: 63 ): “One species of this family, which I obtained here, was new to me: and is certainly very rare: its shell was yellow: rather smaller and more elongated than J. communis [i.e., J. janthina ]; and the whirl more prominent and spiral. The contained animal was also of a yellow colour: but in the form of the float and other respects, it closely resembled the ordinary blue shelled species”. The neotype designated here establishes that this name applies to the shorter Recluzia species identified here as Recluzia lutea . Bennett (1834) earlier described a specimen of Janthina captured at sea at 0°14'S 20°07'W , NW of Ascension Island , central Atlantic Ocean, but did not mention the “yellow species”. Petit de la Saussaye (1853: 118–119 , pl. 5, figs 2–3) named two short-spired species of Recluzia , distinguishing the two by R. rollandiana having more inflated whorls, a more deeply impressed suture, and more pronounced spiral striae than R. jehennei . However, his illustrated shells are both well within the range of variation of R. lutea . A. Adams (1861) provided no dimensions or illustration of Recluzia bensoni , but the holotype was illustrated by Yen (1942 : pl. 19, fig. 122). The illustration and examination of the holotype (NHMUK1878.1.28.394) demonstrate that this is a very small, narrow specimen of R. lutea . Souverbie’s (1872 : pl. 1, fig. 8) illustration of a syntype and examination of the syntypes of R. montrouzieri demonstrate that this also is indistinguishable from the holotype of R. rollandiana , and falls within the variation of R. lutea . Wattebled’s (1886 : pl. 3, fig. 4) illustration of a syntype and examination of the syntypes of Recluzia rollandiana var. annamitica show that this also is closely similar to the type material of R. rollandiana and R. jehennei , and again is part of the variation of R. lutea . Four specimens ( Figs 36M–P ) from the same sample from Great Exhibition Bay, Northland , New Zealand , are illustrated to show the range of variation of R. lutea , although they do not include the most extreme specimens observed. The specimen in Fig. 2S–T is from the same sample. A collection of specimens of Recluzia lutea cast ashore on the beach at Taupiri Bay, Northland , New Zealand (NMNZ M081518, collected by A. Allo, 24 Mar 1982 ; 12 moderate-sized to large shells plus 14 small juveniles) consists of specimens ranging in height from 29.4 mm to larval shells. Some of these were studied by SEM ( Figs 37A–G ). All specimens less than c. 13 mm high are short and wide and closely resemble Alexania natalensis and the holotype of R. globosa in all characters. Several have dried floats attached, each now forming a thin, dark brown, rigid, irregular mass. Several larval shells were incorporated in the float material of some of the moderate-sized specimens (some now separated). One small shell has 10 larval shells attached to the exterior of its teleoconch by dried float material ( Figs 37C–D, G ). These confirm statements by Colman (1986) and Churchill et al . (2011a) that juveniles of R. lutea , including recently metamorphosed larval shells, live attached to the floats of adults. Also, a photograph of a living specimen of R. lutea published on a web page ( Churchill et al ., 2011b : figs 1A–B; Riek, 2017 ; Fig. 6C ) clearly shows a relatively large juvenile specimen of R. lutea on the float of an adult female, lying between the float and the foot. The protoconch in the specimens studied by SEM, although very similar to that of Janthina species, with very similar sculpture and an identical protoconch 1, is shorter and has a protoconch 2 of only 2.2 whorls, rather than 3.2 whorls in the Janthina species available for study. However, the significance of this apparent distinction is uncertain, in view of the lack of knowledge of the number of protoconch whorls in J. pallida .