Beu: Evolution of Janthina and Recluzia 
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other necessities of the neustonic habit. The minor shape and 
sculptural characters used by, e.g., Morch (1860) and Iredale 
(1929) to distinguish genera or subgenera within Janthina 
are regarded here as merely species characters. The groups 
segregated by Morch (1860) were treated as sections of 
Janthina by Thiele (1929: 225), but have been used by few 
other authors. Whether the adult lays egg capsules or broods 
its eggs in the oviduct also is a developmental difference, of 
no phylogenetic significance (Bouchet, 1990). 
Included species. Species included here in Janthina 
are listed and revised in their order of appearance in the 
stratigraphical record. 
1 Janthina typica (Bronn, 1861) (= Janthina hartungi Mayer, 
1864, Heligmope dennanti Tate, 1893, Turbo postulatus 
Bartrum, 1919, Acrybia (Hartungia) chouberti Chavan, 
1951, Hartungia elegans Tomida & Itoigawa, 2001, 
Eunaticina abyssalis Simone, 2014), late Miocene-early 
late Pliocene (Messinian-early Piacenzian), Azores, 
Madeira, Selvagem Is., SW Atlantic, Morocco, New 
Zealand, southern Australia, Japan; presumably formerly 
cosmopolitan. 
2 Janthina krejcii sp. nov., Pliocene (Zanclean), Santa Maria 
I., Azores; presumably fonnerly cosmopolitan. 
3 Janthina chavani (Ludbrook, 1978) (= Parajanthina 
japonica Tomida & Itoigawa, 1982, Kaneconcha knorri 
Kaim, Tucholke & Waren, 2012), late Pliocene-early 
Pleistocene (late Piacenzian-Calabrian), southern 
Australia, New Zealand, Japan, mid-Atlantic ridge; 
presumably formerly cosmopolitan. 
4 Janthina globosa (Swainson, Jan 1822) (= J. prolongata 
Blainville, Aug 1822), Piacenzian/Gelasian-present day, 
cosmopolitan in warm seas; Piacenzian-Gelasian fossils 
from Jamaica and Luzon, Philippines. 
5 Janthina janthina (Linnaeus, 1758) (30 synonyms), one 
Holocene fossil in New Zealand, many reported from 
core tops in the Mediterranean Sea and the Cariaco Basin, 
Caribbean; living, cosmopolitan in warm seas. 
6 Janthina exigua Lamarck, 1816, living only, cosmopolitan 
in warm seas. 
7 Janthina pallida Thomson, 1840, living only, almost 
cosmopolitan in warm seas; not recorded from New 
Zealand or eastern Australia. 
8 Janthina umbilicata d’Orbigny, 1841, living only, 
cosmopolitan in warm seas. 
Taxa not included. The following taxa have been included 
in Janthinidae by earlier authors, but are excluded here from 
Epitonioidea: 
1 Roding (1798: 75-76) included six species in Janthina , 
first section, “Elevata”, and a further nine species and four 
varieties in a second section, “Depressiuscula”. The second 
section obviously has nothing to do with Janthina as used 
now, as most names refer to Gmelin (i.e., mostly Linnaeus) 
Helix species (H. pomatia, with two varieties; H. scalaris, 
H. ligata, H. jamaicensis, H. picta ) and three are nomina 
nuda. Four of the six names in section Elevata also are 
nomina nuda , none of which has been referred to again 
to the writer’s knowledge (J. singidaris, J. turbinoidea, J. 
limbata, J. pellucida ). The fifth name, J. cytherea, refers 
to Chemnitz (1786: pi. 123, fig. 1063), a basal view of a 
small blue-grey shell on a plate showing terrestrial snails, 
so this is an available but probably unidentifiable name. 
2 Janthina alba Anton, 1838: Anton (1838: 50) noted that 
the specimen he described as Janthina alba might really 
be a young specimen of Helix pisana. Dr K. Schniebs 
(Senckenberg Naturhistorische Sammlung Dresden, pers. 
comm. 23 Feb 2016) stated that Anton’s own copy of his 
catalogue has a hand-written note by Anton: “It is Helix 
pisana Mllr. juv”, so this name is a synonym of Theba 
pisana (Muller, 1774), Helicidae. A type specimen is not 
present in Dresden. 
3 Kaiparathina Laws, 1941: Laws (1941) presumably 
composed this generic name because he thought the shallow 
sinus in the outer lip of the early Miocene type species 
indicated that it is related to Janthina ( K. praecellens 
Laws, 1941, Pakaurangi Point, Kaipara Harbour, New 
Zealand; Otaian New Zealand Stage, 21.7-18.7 Ma, 
late Aquitanian-early Burdigalian). He did not refer the 
genus to a family, apparently deliberately, although he 
compared it with some of the characters of Heligmope (i.e., 
Hartungia ; synonymized here with Janthina). This small 
(height 3.2 mm), thick, aragonitic shell with a nacreous 
inner layer was referred to the Trochidae by Beu (1973) 
because a nacreous inner shell layer is plesiomorphic for 
Vetigastropoda, particularly Trochoidea and Turbinoidea 
(Williams et al., 2010). This position was confirmed by 
Marshall (1993) when he described five living Southwest 
Pacific species, recorded fossils as old as early Eocene, and 
referred the genus to Margaritinae Kaiparathini. Williams 
(2012: 589) placed Kaiparathina in family Trochidae, 
subfamily Kaiparathininae Marshall, 1993. 
4 Edithais pehuensis (Marwick, 1926: 319, pi. 73, figs 6, 
8), late Miocene (Tongaporutuan New Zealand Stage, 
Tortonian); referred to Lippistes Montfort, 1810 by 
Marwick (1926), to Heligmope by Finlay (1931: 5) 
following a suggestion by J. Marwick, and to Hartungia by 
Fleming (1953a; 1966:49, pi. 90, figs 948,950), Tomida & 
Nakamura (1981) and Tomida & Kitao (2002). However, 
it was referred to Concholepas Lamarck, 1801 by Beu 
(1970) and to the cosmopolitan muricid genus Edithais 
Vermeij, 1998 by Vermeij (1998). This unusual shell with 
an isostrophic spire and regular, low, wide spiral cords has 
a general resemblance to Janthina typica , but the enlarged 
spiral ridge bordering the wide umbilical hollow fades out 
before the aperture, i.e., it is not generated by a sinus in the 
lip, the axial ridges are low, wide and widely spaced, and 
the spire is below the top of the aperture. The similarity is 
increased by its thin, dark brown, calcitic outer layer over a 
thicker aragonitic inner layer, but this is seen also in many 
other muricids. Holotype (still the only known specimen): 
GNS TM4494, mid-Tongaporutuan New Zealand Stage 
(mid-Tortonian, late Miocene, c. 9 Ma), Okolce Road, 
1.2 km west of Pehu trig, station, Waitara district, North 
Taranaki. Edithais , Family Muricidae. 
5 Janthina cimbrica Sorgenfrei (1958: 176, pi. 32, fig. 110), 
Arnum Formation, Denmark (Miocene), from a depth of 
55 m in a well at Glejbjerg. This minute (H 1.68, D 1.37 
mm) evenly inflated shell with sculpture of microscopic 
cancellate lirae does not belong in Epitoniidae. It is a 
larval shell, type species of Mioseguenzia Nordsieck 
(1973) {nomen nudum), recognized as cypraeacean 
by Quinn (1983: 727, 744) following comments by P. 
Bouchet, and referred to Cypraea by Janssen (1984: 195). 
Janssen (1984: pi. 8, figs 7-8) illustrated very similar 
protoconchs of Cypraeidae from Winterswijk-Miste, the 
