Contents

Foreword
Introduction
Geography in this publication (Figure 1)
Rejected Records not included in this publication
Abbreviations used
Taxonomy
Genus Acirsa Mörch, 1857
-Subgenus Acirsa s.s
-Subgenus Plesioacirsa De Boury, 1909
Genus Kurodacirsa Masahito & Habe, 1975
-Subgenus Kurodacirsa s.s
-Subgenus Minabescala Nakayama, 1994
Genus Alexania Strand, 1928
Genus Alora H. Adams, 1861
Genus Amaea H. & A. Adams, 1853
-Subgenus Amaea s.s
-Subgenus Narvaliscala Iredale, 1936
-Subgenus Filiscala De Boury, 1911
-Subgenus Acrilla H. Adams, 1860
-Subgenus Scalina Conrad, 1865
-Subgenus Clathroscala De Boury, 1889
Genus Cirsotrema Mörch, 1852
-Subgenus Cirsotrema s.s.
-Subgenus Elegantiscala De Boury, 1910
-Subgenus Discoscala Sacco, 1891
-Subgenus Boreoscala Kobelt, 1902
Genus Claviscala De Boury, 1909
Genus Compressiscala Masahito & Habe, 1976
Genus Constantia A. Adams, 1860
Genus Cycloscala Dall, 1889
Genus Eccliseogyra Dall, 1892
Genus Eglisia Gray, 1847
Genus Epitonium Röding, 1798
-Subgenus Epitonium Röding, 1798 s.s.
-Subgenus Mazescala Iredale, 1936
-Subgenus Kiiscala Nakayama, 1995
-Subgenus Lamelliscala De Boury, 1909
-Subgenus Hirtoscala Monterosato, 1890
-Subgenus Hyaloscala De Boury, 1889
-Subgenus Laeviscala De Boury, 1909
-Subgenus Nitidiscala De Boury, 1909
-Subgenus Depressiscala De Boury, 1909
-Subgenus Parviscala De Boury, 1887
-Subgenus Perspicioscala Nakayama, 2000
-Subgenus Asperiscala De Boury, 1909
-Subgenus Pupiscala Masahito, Kuroda & Habe, 1971
-Subgenus Papyriscala De Boury, 1909
-Subgenus Limiscala De Boury, 1909
-Subgenus Foliaceiscala De Boury, 1912
-Subgenus Globiscala De Boury, 1909
Genus Gyroscala De Boury, 1887
-Subgenus Pomiscala Iredale, 1936
-Subgenus Circuloscala De Boury, 1887
-Subgenus Fragiliscala Azuma, 1972
Genus Obstopalia Iredale, 1936
Genus Opalia H. & A. Adams, 1853
-Subgenus Nodiscala De Boury, 1889
-Subgenus Pliciscala De Boury, 1887
Genus Opaliopsis Thiele, 1928
Genus Papuliscala De Boury, 1991
Genus Periapta Bouchet & Warén, 1986
Genus Plastiscala Iredale, 1936
Genus Rutelliscala Kilburn, 1985
Genus Sagamiscala Masahito, Kuroda & Habe in Kuroda, Oyama & Habe, 1971
Acknowledgements
References
Index
Colour Plates 1-20

Foreword

Studying Epitoniidae is very laborious work due to their nature. Most of the species in this family are ‘rare’, so that only small numbers of individuals are available for study (usually from private collections).
Therefore, many collectors listed in the acknowledgements are collaborating to this study. After gathering many data on the specimens and references, some great misunderstandings or misidentifications were corrected, and some new species were described to complete this study. However, due to some urgent project on Japanese molluscs by Okutani (2000), the new species were unfortunately reported to ‘Venus’ shell magazine in 2000, previous to the publication of this book.
This study is based on the review of North Pacific Epitoniids in 1996-1998 and includes some minor changes since then. One new species is introduced in this work.
July, 2003
Taisei Nakayama, Ph.D.

Introduction

The Epitoniidae are one of the most complicated families among gastropods, usually classified as heterogastropods, and occurring in all seas from the lower intertidal zone to abyssal seabeds. As far as the biology is known, all species are associated with coelenterates, either as ectoparasites or as foraging predators. Since the Linnaean era, many species have been reported and described as new, but the actual knowledge of the taxonomy is rather chaotic. Some systematic reviews have been done by Sowerby (1844, 1873), Tryon (1887), Clessin (1897), and De Boury (1889) on world-wide species, by Clench & Turner (1950), DuShane (1974), and Bouchet & Warén (1986) on north Atlantic species, and by Kilburn (1985) on South African species, but few revisions have been made of Western Pacific species. Shikama & Horikoshi (1963), Shikama (1964), and Matsukuma et al. (1991) figured epitoniids from Japan and some other regions in illustrated mollusc publications, but did not provide any detailed descriptions. Kira (1959) and Habe (1961) illustrated some epitoniids in their books on Japanese molluscs; most of the listed species are common and generic placements are not discussed. After that, Kuroda, Habe & Oyama (1971) studied the molluscs from Sagami Bay (Pacific coast of central Japan) and illustrated and described some new minute epitoniids. Kay (1979) illustrated and described some new epitoniids from Hawaii. In the publications of Springsteen & Leobrera (1986) from the Philippines, Powell (1979) from New Zealand, and Wilson (1993) from Australia, several epitoniids are illustrated.

De Boury (1909) proposed a lot of subgenera in the Epitoniidae, and most of them are used in later studies as full genera without precise discussion. This is caused by the insufficient descriptions of De Boury on the one hand and the conchological trend of expanding generic classification on the other hand. Thiele (1928), Clench & Turner (1952), and Azuma (1972) tried to use radula formula’s for classification, but radulae of epitoniids show large individual variation. Taki (1956, 1957) studied the anatomy of epitoniids, but these studies did not help much for the classification either. Kilburn (1985) pointed out that radula studies are useless for generic classification, whereas protoconch morphology gives important information for classification and is thus crucial in solving the generic chaos. Of course shell characters are important as well: the presence or absence of a basal cord, axial ribs or lamellae, spiral ribs or grooves and suture characteristics are major points to classify the genera.

There are many indistinct species which were described without any illustrations in the 19th century by e.g. A. Adams (1861, 1863). Sowerby (1844, 1873), Tryon (1887) and Clessin (1897) illustrated some minute species with poor figures, which are also ambiguous. Kuroda & Habe (1952), and Higo & Goto (1993) listed the Japanese molluscs, in which the status of these obscure species is not clear. Moreover, Kuroda & Habe (1954), and Fukuda et al. (1990) tried to clear these species, but the descriptions are somewhat different from the deposited Adams specimens in the Redpath Museum of McGill University in Canada. Kaicher (1980-3) solved a number of these ambiguous species by publishing photographs of type specimens preserved in the Natural History Museum, London (BMNH) or in the Museum national d’Histoire naturelle in Paris. Moreover, Cernohorsky (1972) and Kilburn (1985) also illustrated some of Sowerby’s type specimens in the BMNH. Those works helped to recognise some ambiguous species listed by Higo & Goto (1993) and Kuroda & Habe (1952).

Contrary to many recent species, fossil species are not studied sufficiently in Japan. Yokoyama (1922, 1926, 1927) studied Pliocene fossils from the Kwanto region (around Tokyo) and Ozaki (1958) studied the same area. They reported some new fossil epitoniids. Some of them also occur as recent species. In this work, some of them are studied for comparison with recent species. Bouchet (1997) pointed out that the taxonomic studies in Mollusca are in chaos and many scientific names are synonymised by systematic studies. The major purpose of this study is to solve part of the taxonomic chaos in this family, based on species collected around Japan and the adjacent area in the North Pacific.