logical Journal o f t h e Linnean Sociely (1990), 100: 1-25. With 3 figures
Species of Naticidae (Mollusca: Gastropoda)
described by Linnaeus in the Systema Naturae
( 1758)
ALAN R. KABAT
Museum
of Comparative <oology,
Harvard University, Cambridge, M A 02138 U.S.A.
Rcceived March 1989, accepted for publication December 1989
In the Systema Naturae (1 758) Linnaeus described seven species now placed in thc gastropod family
Naticidae. This study documents their identity based on critical re-examination of original
descriptions and original specimens in the Linnean Society of London and the Zoologiska Museet of
Uppsala Universitet. The six valid species are redescribed; the other is a nomen dubium. Four of the
species are type species of currently accepted naticid genera.
KEY WORDS:- Linnaeus
-
Naticidae
-
Gastropoda
-
Prosobranchia.
CONTENTS
Introduction . . . . . . . .
Methods . . . . . . . . .
Classification of the Naticidae . . .
Genus Sinum Roding, 1798
. . .
Genus Naticu Scopoli, 1777
. . .
Genus Naticnriur DumPril, 1806 . .
Genus Polinices Montfort, 1810
. .
Genus Neuerita Risso, 1826
. . .
Nerita glaucina Linnaeus, 1758 (nomen dubium)
Acknowledgements
. . . . . .
References
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INTRODUCTION
I n the 10th and 12th editions of the *sterna Naturae, Linnaeus described seven
species of gastropods which were subsequently placed in the Naticidae
(Prosobranchia, Mesogastropoda) (Linnaeus, 1758, 1767). Two were originally
described in Helix, and five in Nerita. The former are now placed in Sinum
Roding, 1798 and the latter in Natica Scopoli, 1777, Naticarius Dumtril, 1806,
Polinices Montfort, 1810, and Neverita Risso, 1826. Of the seven naticid species of
Linnaeus, four are type species of various naticid genera. Currently, only three of
Linnaeus’ naticid species are clearly defined; the other four have often been
misrepresented or ignored. The purpose of this paper is to document the correct
identity for these seven species in order to resolve long-standing problems in the
Present address: Division of Mollusks NHB-118, National Museum of Natural History, Washington D.C.
20560 U.S.A.
1
0024-4082/90/090001+ 25 $03.00/0
0 1990 The Linnean Society of London
2
A. R. KABAT
taxonomy of the Naticidae. T h e valid species are redescribed and the junior
synonyms arc indicated. This paper is based on examination of the type material
in the Linnean Society of London (LSLj and the Zoologiska Museet, Uppsala
Lniversitet (ZMUC‘) arid on a re-evaluation of the original descriptions and the
references cited. T h e type material is clearly delimited and lectotypes are
selected. as appropriate. T h e geographical range of each species is indicated but
specific type-localities are not designated as it is not possible to restrict the type
material in this manner.
First, i t is necessary to consider the history of Linnaeus’ works and the
vicissitudes of his shell collections in order to properly analyse his species.
Perhaps the most important consideration is that Linnaeus himself did relatively
little travelling or collecting outside Sweden, and hence much of his shell
collection was acquired from other collectors, who were not always concerned
with providing accurate locality information. In addition, Linnaeus worked on
the royal conchological cabinets of Sweden (i.e. the ‘Museum Ludovicae
Ulricae’i; these shells are also often of doubtful provenance. After his death the
Linnaean collection did not remain inviolate, but was subject to a number of
unfortunate events which did not increase its value. His personal collection was
cvcntually acquired by the Linnean Society of London, and is now kept at
Burlington House after having been moved at least four times (Gage & Stearn,
1988i. FVhilc that collection was still in Sweden and shortly after its arrival in
London, various well-meaning persons undertook b add specimens in order to
fill presumed gaps or to otherwise provide new material (i.e. non-Linnaean
species;, as discussed by Dodge (195913: 175) and Dance (1967: 3-4).
The first conchologist to kvork critically on this material was Sylvanus Hanley
who attempted to analyse all of the molluscan species of Linnaeus, based on the
I S L holdings (Hanley, 1855). Supposedly, he did not rearrange the collections
at t h a t time; nevertheless, he (and perhaps others) subsequently attempted to
reorder the shells and did not always follow careful curatorial procedures
[Dance, 1967: 5).Jackson (1913) published a catalogue of the specimens present
but he did not differentiate between the Linnaean and the non-Linnaean
material. Dodge (1952--1959a) wrote a series of papers on various Linnaean
species (not including naticids j based primarily on a n uncritical acceptance of
Hanley i1855). Finally, Dance ( 1967) thoroughly restudied the LSL material
and critically evaluated the various lots to determine whether or not they were
truly Linnaean in origin.
‘ l h e unfortunate conclusion is that one can rarely be absolutely certain that a
given specimen is the actual Linnaean specimen since the original descriptions
are sufficiently broad that almost any conspecific shell can appear to ‘fit’ the
description. Hence, the presence of a certain shell which corresponds to the
description does not prove that this specimen actually belonged to Linnaeus.
Dance i 1967: 20-~22)listed those Linnaean species which were not present in the
LSL collections; however, his lengthy manuscript on those species which are
present remains unpublished. There is also a ‘residual collection,’ temporarily
housed in the British Museum (Natural History) which contains various shells
that probably are not original Linnaean specimens, although some represent
1,innaean species. This latter collection is of no further value in establishing the
Linnaean naticids due to its dubious provenance.
The second Linnaean collection, and certainly the less-appreciated, is that of
LINNAEUS’ NATICIDAE
3
the ‘Museum Ludovicae Ulricae’ now housed in the ZMUU. Most zoologists are
aware that Linnaeus’ 10th and 12th editions of his Systema Naturae form the
foundation for modern taxonomy, but few are also cognisant of his
contemporaneous work, the catalogue of the Swedish royal natural history
collections, known as the Museum Ludovicae Ulricae (M.L.U.) (Linnaeus, 1 764).
Although written in the early 175Os, its publication was delayed until after the
appearance of the 10th edition of the Systema Naturae (Dodge, 1959b: 176). The
M.L. U. is of importance as it provides supplementary diagnoses and references
for many (but not all) species described in the Systema Naturae. Correlated with
the M.L.U. were the collections of the Royal Family; again, like Linnaeus’ own
collections, these have not remained inviolate over the years. Even though the
royal collection has remained in Sweden, it has suffered from some rearranging,
but not as much from the addition of non-Linnaean material. Hanley (1859)
compared Linnaeus’ original manuscript of the M.L. U. with the published
version, and compiled an annotated checklist of the discrepancies in the species
descriptions. Lovtn (1887),provided a detailed history of the M.L.U., noted that
Linnaeus had not labelled the specimens, bewailed those authors who had
ignored this collection and its catalogue, and was perhaps overly optimistic in his
view that although the collections had been subsequently labelled (in part) by a
botanist, they had not suffered from any improper handling. Holm (1957)
further reviewed the history of the M.L.U. collections, as well as that of several
lesser collections in Uppsala. Odhner ( 1953, unpublished) provided a complete
catalogue of the Linnaean shells in the ZMUU, along with his remarks on their
current systematic status.
Thus, we are left with two independent collections of ‘Linnaean shells’, both of
which have been subject to rearrangement and potential adulteration. As Dodge
(1959b) and others have emphasized, a number of aspects must be considered
when analysing the Linnaean species. First, the original description (diagnosis)
of the species: while this is normally the basis for modern nomenclature, the
descriptions written by Linnaeus are sometimes too brief or too vague to
designate a single modern species. Second, the references cited by Linnaeus are
of extreme importance since Linnaeus did not provide his own illustrations, but
merely referred to previous iconographies (Boss, 1988). Again, however, these
illustrations often suffer from a combination of poor printing (resulting in
indeterminate identifications) and a rather broad species concept. The
illustrations cited for a single Linnaean species may include several species.
Third, the localities cited by Linnaeus are unreliable, being too broad or even
completely erroneous, since Linnaeus often had to rely on the locality
information provided by amateur collectors. Finally, there are the collections of
Linnaeus, now housed in London and Uppsala. Their condition and veracity has
been discussed above. It will be seen that the analysis presented herein of the
Linnaean species of Naticidae must be based on a critical evaluation of the above
factors, combined with our modern knowledge of these species.
Several previous studies on specific groups of marine molluscs in the Linnaean
collections should be mentioned, since they serve as a guide to further
malacological work. First, Dodge (1952-1959a) is not of great value since it is
based solely on a microfilm of the specimens in the LSL. Schilder (1966) studied
the cowries (Cypraea); Olsson & Dance (1966) the Olividae; and Nelson & Pain
(1986) the whelk Neptunea. These three papers are all based on material in the
A. R . KAB.Xl
4
LSL. Kohn 1963) made a comprehensive study of the cone shells (Conus) based
on a thorough consideration of the specimens in the LSL and ZMUU.
I
LIE 1 HODS
’ l h e references cited by Linnaeus and subsequent authors were consulted to
determine the species that were represented. S) non).mies for each species are
prcsented, including the junior synonyms and references to the major 19th and
20th-ccntur) molluscan iconographies and regional monographs.
-1hc geographical ranges for the naticid species recorded in this paper are
bawd on examination of the mollusc collections in the Australian Museum,
S>dne) ; M I S j , Bernice Pauahi Bishop Museum, Honolulu (BPBM), Museum
of Comparative Zoology, Hari ard University IMCZ) and Western Australian
lluseum, Perth ‘II’All,.
?‘hc Xaticidac is a cosmopolitan family of predatory marine gastropods which
burrow in soft-bottom sediments. T h e family can be divided into four subfamilies
,~llarincovich,1977J ; representative genera are indicated: ( 1 ) Ampullospirinae,
a mostly extinct assemblage of high spired shells with tabulate or canaliculate
whorls :A,lrnnuropsis, Globularia); (2) Polinicinae. wide-spread, with usually
monochroinatic shells (Polinices, Euspirn, u’lererila); ( 3 ) Sininae, tropical, with low
spired to auriforni striated shells (Sinurn, Eunaticina); and ( 4 ) Naticinae, mostly
tropical, with calcareous opercula and often multicoloured shells (Natica,
.Yuticnrius, C‘cyptona&ica).However, with over 200 described genera and almost
3000 described species [Triassic to Recent ), the lower level taxonomy of this
family requires considerable further research.
Genus Sinurn Roding, 1798
Rijding 1798: 14) established the genus Sinurn for two species of naticid
gastropods: Sinurn fuscum Roding, 1798 and Helix haliotoidea “Gmel.”. T h e generic
description was singularly brief “Milch-Napchen”. The first species, although
based on some of the same illustrations that Born ( 1 778) and Gmelin (1791) had
used for Helix haliotoidea, is not directly referable to any one species o f S i n u m and
is now a nornen dubium. The second species was originally described by Linnaeus
(17581, and its status will be discussed later in this paper. Roding’s work was
overlooked during much of the 19th century, as Lamarck’s 1799 masterpiece,
“Prodrome d’une nouvelle classification des coquilles . . .” was long used as a
basis for molluscan generic nomenclature. I n that work, Lamarck (1799: 77)
described the genus Sigarelus, with a single included species, Helix haliotoidea
Linnaeus, 1758. Subsequently, Dall ( 1906) resurrected Roding’s work and
demonstrated that many of the taxa described by Lamarck and other
contemporaries were actually junior synonyms of Roding’s taxa. Thus, Dall
( 1906: 294) placed Sigarelzis Lamarck, 1799 in the synonymy of Sinurn Roding,
1798 and later (19 15: 109) designated the type of Sinurn as Helix haliotoidea
I.innaeus, 1758 ii.e. the second species listed by Roding). Both species in
LINNAEUS’ NATICIDAE
5
Roding’s Sinum have been generally ignored in this century except that haliotoidea
has been cited as the type species (4.Majima, 1989: 69-70).
Sinum neritoideum (Linnaeus, 1758)
(Fig. lA, B)
Synorymy
Helix neritoidea Linnaeus, 1758: 775, No. 619 [Cited from M.L.U.; Gualtieri
(1742), pl. 64, fig. I. Locality: unknown].
Helix neritoidea Linnaeus, 175%. Linnaeus, 1764: 672, No. 380; Linnaeus,
1767: 1250, No. 711 [added citation of “Mus. L. U. 672, n. 380”]; Gmelin,
1791: 3663; Turton, 1802; 540; Hanley, 1855: 390, 541.
Sigaretus neritoideus (Linnaeus, 1758). Reeve, 1864: pl. 1, figs 5a, 5b; Sowerby,
1882: 40, pl. 441, fig. 1; pl. 442, figs 16, 17; Weinkauff, 1883: 18-20, pl. 3,
figs 7-1 1; Tryon, 1886: 55, pl. 22, fig. 35; pl. 23, figs 38-40.
Non “Helix neritoidea Linnaeus, 1758”. Dillwyn, 1817: 972 [“Inhabits the
Baltic”; probably a lamellariid].
Non “Sigaretus neritoideus Linnaeus, 1758”. Rkcluz, 1843: pl. 1, fig. 7; pl. 2,
fig. 3; pl. 4, fig. 11 [Not described in the text; illustrations may be Sinum cymba
(Menke, 1828)l.
Description
Shell oval (shell axis to 35 mm; breadth to 43 mm; width to 38 mm),
moderately convex and thickened, with well-developed spiral ridges extending
across the entire last whorl. Protoconch and early teleoconch distinctive bluishpurple; later teleoconch off-white with pale yellowish periostracum (often faded
or eroded). Sporadic irregular concentric growth lines extending across last
whorl resulting in slight interruption of spiral ridges. Suture sharply demarcated
but shallow, adpressed. Aperture nearly circular, strongly prosocline. Shell
interior smooth, slightly darkened. Peristome simple; outer lip thick; inner
(columellar) lip well developed with the columella reflected slightly over the
preceding whorl. Shallow umbilical region, closed off adapically by the parietal
callus (inducturum). Among the largest-sized species in the genus.
Material examined
The type material in the LSL consists of one box, No. 560, containing four
conspecific shells, all referable to Sinum neritoideum. They are higher spired than
S. haliotoideum and have the distinctive purple larval shell. The ‘type material’ in
the ZMUU consists of one tray, No. 380, containing one shell, which, however, is
actually a Stomatella (Trochidae). Odhner (1953: 19) suggested that this is
Stomalella papyracea “Chemnitz” (Gmelin, 1791). I t is obvious that some mixing
of lots or the introduction of non-Linnaean material has occurred here. The
third-largest shell in the LSL, Box No. 560 is here selected as the lectotype of Helix
neritoidea Linnaeus, 1758 (Fig. 1A). The remaining three specimens are
paralectotypes (Fig. 1B) .
Remarks
The cited figure from Gualtieri (1 742: pl. 64, fig. I) is not referable to any one
species of Sinum. Sinum neritoideum (Linnaeus, 1758) is similar to S.javanicum
6
.I R KARAT
cGri&th & Pidgeon, 1834) and S. laei*igatum (Lamarck, 1822). The former is
somewhat thicker, more convex and with broader ridges than S. nerilozdeum; thc
latter is higher spired and has a distinctive brownish-red interior. All threc
species occur in the central and western Indo-Pacific.
Geographical iange
This rare species has been found in the central portions of the tropical
Indo-Pacific: Sagami Bay and Kumano Sea, Japan southwards through
Singapore, Java, P a p a Netv Guinea and New Caledonia to Queensland,
Australia.
Sinum halzotozdeum (Linnaeus, 1758)
(Fig. 1C-E)
S yn onznz_y
Helzr halzotozdea Linnaeus, 1758: 775, No. 62 1. [Cited from Petiver ( 1702-06),
pl. 12, fig. 4; Rumphius ( 1 705), pl. 40, fig. R; Gualtieri ( 1 742), pl. 69, fig. F;
.lrgen\ i l k 1 1742), “pl. 8, fig. D” {error for pl. 7, fig. C; see Emendanda,
Linnaeus, 1758: 824). Locality given as “M.Mediterraneo, Arnericano”.]
Hdrx haliotoLdea Linnaeus, 1758. Linnaeus, 1764: 673, No. 382. [Cited from:
Rumphius i1705), Gualtieri ( 1742), and Argenville ( 1742); the Petiver ( 1 70206, reference was omitted.];
Linnaeus, 1767: 1250 1, No. 713 [four
additional illustrations: Lister ( 1685 92), pl. 570, fig. 21; Buonanni (1709),
pl. 475, fig. 404; Klein ( 1753), pl. 7, fig. 1 14; and Adanson ( 1 757), pl. 2, fig. 2.
Imcality: *‘M. Mediterraneo, Arnericano, Asiatiro”.]; Born, 1778:
408 409; Turton, 1802: 540-1.
Helix helzotozdea [sic] (Linnaeus. 17583. Gmelin, 1791: 3663 4.
Szgarefur helzotoideus [sic] (Linnaeus, 1758). Bosc, 180 1: 255.
SzgmetuJ halzotozdeus (Linnaeus, 1758). Lamarck, 1822: 208; Deshayes, 1832:
949--950; Deshayes, 1843: 9 10; Sowerby, 1882: 42-43, pl. 441, fig. 5,
pl. 442, fig. 3 1.
Cryplustoma halzotozdeum (Linnaeus, 1758). Gray, 1826: 49 1.
Sigareiur pfanatuJ (Rkcluz, 1843: 1 , 6 ) . [Cited from Gualtieri, 1742, pl. 69, fig. F;
type locality (restricted by Kilburn, 1976: 874) Maht, Seychelles. Syntypes,
hfuseuni d’Histoire Naturelle de Gen6L.e (No. 1 152/30); examined.]
‘Czgaretur planzilatus Rtcluz, 1813: pl. 3, fig. 4; Rkcluz, 1844a: 21 [a variant
>pelling of the preceding]; Reexe, 1864, pl. 2, figs 7a, 7b; -Sowerby, 1882:
42, pl. 442, figs 29, 30; Ll’einkauff, 1883: 15 16, pl. 2, figs 7-9; pl. 4,
hgs 10-12; Tryon, 1886: 58, pl 22, figs 75-77.
Helix halzotzdea [sic] Linnaeus, 1758. Hanley, 1855: 390 1 , 541, pl. 4, fig. 7.
[Hanley noted that thr cited references of Linnaeus “are almost too rude to
admit of incontestable recognition” and believed this to be a tropical species.]
Szgaretus m b e r i Bartsch, 1918: 187. Not illustratrd. Palawan, Philippines.
Holotype, USNM 2 19050, examined.
Sznum planatum (RCcluz, 1843). Kilburn, 1976: 874-6, figs 23-24 [and references
therein; exclu$i\Teof Ectosznum paidocont‘exum Iredale, 1931 = Sznum zonale (Quoy
& Gairnard, 1832)l.
.4on “Helzx halzotozdea Linnaeus, 1758”. Dillwyn, 1817: 973 [may be a
lamellariid gastropod].
LINNAEUS’ NATI C IDAE
Figure I . A, Sinum nerztoideum (Linnaeus, 1758). Lectotype, LSL (No. 560), apertural view. Shell axis
25.3 mm; breadth 32.5 mm, width 25.6 mm. B, Sinum nel-itoideum (Linnaeus, 1758). Paralectotype ( 1
of 3), LSL (No. 560), apical view. Shell axis 27.2 mm; breadth 36.7 mm, width 30.6 mm. C, Sinum
halzoloideum (Linnaeus, 1758). Lectotype, ZMUU (No. 382), apertural view. Shell axis 5 mm;
breadth 38.2 mm, width 25.5 mm. D, Sinurn haliotoideum (Linnaeus, 1758). Lectotype, ZMUU
(No. 382), apical view. E, Sinum haliotoideum (Linnaeus, 1758). Paralectotype, ZMUU (No. 382j,
apertural view. Shell axis 5.6 mm; breadth 35.2 mm, width 25.2 mm.
7
8
-4,R. KABAT
Fleming, 1828: 360; Brown,
1827: pl. 44, figs 1 , 2; Brown, 1844: 23, pl. 2, figs 1, 2 [all these references are
of lamellariids].
.%in “Sigaretus haliolideus [sic] (Linnaeus, 1758)”. Philippi, 1844: 144, pl. 1,
fig. 6 [“Mare Mediterraneum”; is Sinum philippi (Weinkauff, 1883)].
-/Voiz “Sigareius haliotoideus (Linnaeus, 1758)”. Reeve, 1864, pl. 1, figs 4a, 4b.
/-“Mouth of the Gambia, West Africa”; is Sinum concauum (Lamarck, 1822).]
;Van ”Signrefus haliotoideus (Linnaeus, 1 758)”.
DPrcripiion
Shell oblong-oval (shell axis to 6 mm; breadth to 40 mm, width to 26 mm),
low-spired, nearly convolute, quite thin, with fine spiral striae (often worndown) extending over the last whorl. Protoconch (small, 2-2.5 whorls) and
teleoconch uniform white, with a thin yellow periostracum (may be eroded).
Distinct fine concentric growth lines evenly spaced across the whorls. Suture
shallow, adpressed, less distinct in the presence of the periostracum. Aperture
elongated, extremely prosocline. Shell interior smooth, glossy white. Peristome
simple; outer lip thin; inner (columellar) lip thin with columellar rim slightly
developed adapically. Small, shallow umbilicus largely covered by the parietal
callus; a distinctive notch may be present (in well-preserved specimens) at the
junction of the inductura and the columellar rim. Vestigial corneous operculum
present.
Material examined
The material in the LSL consists of one box, No. 56 1, containing six planispiral
shells, similar to the illustration in Gualtieri (1742). These shells appear to have
been originally white, although they are now dirty and splotched with dry
mould.
The material in the Z M U U consists of one tray, No. 382, with two shells,
congeneric with the Linnean Society material. The larger shell has a printed
label [“halioloidea”] on the dorsal side. Odhner (1953: 19) claimed that these two
shells are the Western Atlantic Sinum perspectivum (Say, 1831). The shells are
glossy white, have only the finest striae, and are not as low-spired as are the
shells of S. perspectiuum which also have more prominent striae. They agree with
the original description, the figure in Gualtieri ( 1 742) and with the description
by RCcluz of planatus [ =planulatus]. However, they do not represent the same
Sinum species as do the shells in the LSL, which have moderately pronounced
spiral striae across the outer whorl, and a slightly thinner columellar lip. Since
the specimens in the ZMUU, tray No. 382 are best representative of Linnaeus’
concept of haliotoidea they should be considered the type material for this species.
The larger shell (with the printed label) is here designated the lectotype of Helix
halioloidea Linnaeus, 1758 [ =Sinum halioloideum (Linnaeus, 1758)] (Fig. 1C,D);
the smaller shell the paralectotype (Fig. 1E).
Remarks
Not unexpectedly, it appears that several species of Sinum were confounded by
Linnaeus and succeeding conchologists in their presentation of haliotoidea. Of the
illustrations referred to by Linnaeus ( 1758, 1767), only the figure of Gualtieri
‘1742: pi. 69, fig. F) matches the type material. However, Gualtieri actually had
t w o ‘figure F’ on his plate 69; the lower figure agrees with Linnaeus’ description
LI NNAEU S’ NATI C 1DAE
9
of haliotoidea while the upper figure is a dubious representation of a supposed
auriform gastropod. Linnaeus did not specify either figure; it is possible that he
was unaware that there were two. T h e illustration of Adanson (1757: pl. 2,
fig. 2) is that of Sinum concauum (Lamarck, 1822), from West Africa (FischerPiette, 1942: 138-9) (however, Linnaeus did not cite this figure until the 12th
edition). The remaining illustrations (Petiver, 1702-06; Rumphius, 1705;
Argenville, 1742; Lister, 1685-92; Buonanni, 1709; and Klein, 1753) are
indeterminate, generally poor drawings of low-spired Sinum (although Martens
(1902: 124) identified those of Rumphius ( 1705) as “Sigaretus planulatus”).
Rkcluz (1843: 1; 1844: 19, 21) in his discussion of the “planult!es” Sigaretus,
stated that Gualtieri, pl. 69, fig. F “supra” was the cited figure for Linnaeus’
haliotoidea; and he then used Gualtieri, pl. 69, fig. F “inferior” for his new species,
planatus [ =planulatus, a spelling error]. Unfortunately, as stated above, the
original description of Linnaeus (as well as his type specimens) does not
correspond with “fig. F supra” but rather with “fig. F inferior”. There are two
alternative solutions to this nomenclatural problem. One is to rely solely on the
poor illustrations of Gualtieri, use planatus for the common Indo-Pacific species,
and reject haliotoidea as a nomen dubium, in spite of the adequate description and
type material of Linnaeus. This solution would probably necessitate a ruling by
the International Commission on Zoological Nomenclature. The second solution
would be to consider planatus a junior synonym of haliotoidea and to discard
Ricluz’ misguided choice of “fig. F, supra” as the sole representation of
haliotoidea. T h e latter choice is followed herein, since it is mandatory to utilize
Linnaeus’ original description and type material rather than the dubious
speculations in iconographies.
Apparently unaware of the illustrations used by Linnaeus for haliotoidea,
Kilburn (1976: 874-6) in his redescription of Sinum planatum (Rkcluz, 1843), not
only restricted the type locality to “Mahk, Seychelles”, but also noted (after
RCcluz) that the species was based on the “type-figure Gualtieri, 1742, pl. 69,
fig. F, inferior”. Kilburn (1976: 875) stated that “Gualtieri’s shell should be
sought amongst the remnants of his collection now in the Pisa Museum”, but this
is of little relevance since the syntypes of Ricluz’ Sigaretus planatus are in the
Muskum d’Histoire Naturelle de Genkve. Marincovich (1977: 342) stated that
Helix haliotoidea was from “West Africa”; but did not support his conclusion.
Bergh (1907: 108-1 10, pl. 10, figs 4-16) reported on the anatomy of “Sigarelus
planulatus”; although the description is not sufficient to confirm (in retrospect)
the exact species identity.
Geographical range
This species is moderately common in the central and western portions of the
tropical Indo-Pacific: Okinawa, Philippines (Luzon, Bohol, Penang), Malaysia,
Papua New Guinea, Solomon Islands, New Caledonia, Australia (from Keppel
Bay, Queensland to Dampier Archipelago, Western Australia), westwards to
South Africa (Durban).
Genus flatica Scopoli, 1777
Scopoli (1777), in his comprehensive review of the ‘genera’ of organisms and
minerals, provided brief diagnoses for numerous taxa which although
10
A,
R. KABA’I
nomenclaturally genera, are so broad as to correspond to modern families.
Scopoli is tiow credited with the authorship for certain generic names which
were originally published prior to Linnaeus (1758) but were riot used by
Linnaeus. Adanson (17571, in his remarkable work on the fauna of Senegal,
recognized that the traditional group “,l,”erita” actually combined two unrelated
groups of gastropods. One, which he retained as “Yerita”, included herbivorous
gastropods living in rocky areas, and with teeth or ridges on the columellar lip.
’The other group, separated out as “,Vatica”, included predatory gastropods
which burrow in sandy areas, and lack teeth or ridges on the columellar lip.
Linnaeus ( 1 758, 17671, although not using .Xatica, divided the species of his
,,Veritu into three groups: the first (~“urnbilicatae”)corresponds to Adanson’s
“LVa&-a”; arid the following two (“imperforatae”) represent Nerila sensu stricto
(i.e. the modern Xeritidae). I n any case, Scopoli (1777) was the first postLinnaean author to use the name .Vatica “Adanson”. Scopoli’s diagnosis of this
genus was:
.‘Testa unival\,is; umbilicata; apertura subelliptica. Molluscum tentaculis
binis, basi ocellatis, absque columna adiacente, quam hahent
NERI‘I’IDAE. Helix lusitanica, .\‘brila mammillaris, oilellus, albumen, &c. huius
generis” (Scopoli, 1777: 392).
O f the four species (all Linnaean) included by Scopoli, only the last two are
naticids. l‘he first species is a European terrestrial gastropod and the second is
apparently an African freshwater gastropod.
Anton (1838: 31) selected ..\>rita zitellus Linnaeus, 1758 as the type species of
.IpoiiraSropoli, 1777. Several other authors had ‘selected’ Nerita canrena Linnaeus,
17.58 as the type species of ,\hlica; this latter name is not available since it was not
included by Scopoli (contra Lamarck, 1799: 77; Cossmann, 1888: 159; and Dall,
1892: 362, among others). Forbes ( 1838) established the family NATICIDAE,
Lvith the type genus .Lalira Scopoli, 1777.
~ I a t z r nr9zlrllus (Linnaeus, 1758)
(Fig. 2A-D)
.Qn OI2l;??y
Yenla ~11te11zi~
Linnaeus, 1758: 776, KO. 625 [Figure cited: Rumphius (1705),
pl. 22, fig. D. Locality: “0.Asiae”].
C m t n rutelluy Linnaeus, 1758. Linnaeus, 1767: 1252, No. 7 17; Hanley, 1855:
394 5, 541-2.
.”lerzta tufa Born, 1778: 413 414 [Figures cited: Lister (1685-92), 4, pl. 6, fig. 34;
Rumphius ( 1705 1, pl. 22, fig. D; Petiver ( 1 702-06), pl. 1 1, fig. 3; Argenville
17.1.21, pl. 7, fig. Z; Seba (1758), 3, pl. 39, fig. 30; Geve (1755), pl. 27,
fig. 296. Holotype, Naturhistorisches Museum Wien; examined (Fig. 2C) .];
Born, 1780: 398, pl. 17, figs 3, 4: CIvrlin, 1791: 3672; Turton, 1802: 546.
.tenLo leucobonins Gmelin, 170 ’
’ : + i ~ ~ , i tcited:
s
Kammerer, 1786: 187,
pl 12. figs 5, 6. Locality unknu-,:i;.j, I uizon, 1802: 546.
- l l n t a Jpadzcen Gmelin, 1791: 3672 [Figures cited: Chemnitz, (1781), 5, pl. 187,
figs 1872-3; a variety “beta”, from Chemnitz (1781), 5, pl. 188, figs 1896a,b,
1897. Locality: “littora insulae S. Mauritii”.]; Turton, 1802: 546.
LINNAEUS’ NATICIDAE
11
Cochlis albula Roding, 1798: 146 [Figures cited: Chemnitz, 1781, 5, pl. 188, figs
1896a, 189613, 18971.
Cochlis rufescens Roding, 1798: 148 [Figures cited: Chemnitz, 1781, 5, pl. 187,
figs 1872, 18731.
Naticu vitellus (Linnaeus, 1758). BOSC,1801: 288; Deshayes, 1838: 636-7
[Deshayes noted that rufu Born and uitellus were based on the same illustration
in Rumphius (1 705, pl. 22) but incorrectly assumed that Linnaeus had
erroneously cited fig. D instead of fig. A.]; Hedley, 1913: 299-300;
Majima, 1989: 74-7, pl. 10, figs 1-12.
Natica sbadicea (Gmelin, 1 79 1) . BOSC,1801: 289; Dillwyn, 181 7: 980;. Reeve,
1855: pl. 3, figs 9a, 9b.
Natica rufa (Born, 1778). BOX, 1801; 289; Dillwyn, 1817: 980-1; Lamarck,
1822: 201; Deshayes, 1832: 602; Deshayes, 1838: 639-640; Philippi, 1849:
14-15, pl. 2, figs 1, 2; Philippi, 1852: 85, pl. 13, fig. 1; Reeve, 1855: pl. 16,
figs 70a, 70b; Sowerby, 1883: 80, pl. 457, fig. 42.
Natica fuscuta Link, 1807: 140 [Figures cited: Chemnitz, 1781, 5, pl. 188,
figs 1896a, 1896103.
Naticu helvacea Lamarck, 1822: 200 [Figures cited: Chemnitz (1781), 5, pl. 188,
figs 1896a,b, 1897. Locality: inknown. Syntypes, Muskum National d’Histoire
Naturelle de Genkve (No. 1094/60); examined (figured, Mermod, 1953: 1868, fig. 188).]; Deshayes, 1832: 602; Deshayes, 1838: 637.
Nutica globosa “Chemnitz”. Philippi, 1850: 21-22, pl. 3, figs 1, 2; Philippi,
1852: 52, pl. 8, fig. 5; Reeve, 1855: pl. 11, figs 46a, 46b; Sowerby, 1883: 94,
PI. 457, fig. 47.
Natica (Nutica) rufa (Born, 1778). Tryon, 1886: 29-30, pl. 9, fig. 62.
Nutica (Natica) rufa var. spudicea (Gmelin, 1791). Tryon, 1886: 30, pl. 9, fig. 63.
Natica (Natica) helvacea (Lamarck 1822). Tryon, 1886: 30, pl. 9, fig. 64.
Nutica (Natica) vitellus (Linnaeus, 1758). Cernohorsky, 1971: 173-1 76, figs 2-5.
Nerita rufu [sic] Born, 1778. Majima, 1989: 75, 155.
Non “Nerita uitellus Linnaeus, 1758”. Born, 1778: 414-415; Born, 1780:
398; Gmelin, 1791: 3671; Turton, 1802: 545 [Is Natica stellala Hedley,
19131.
Non “Nerita rufa Born, 1778”. Montagu, 1808: 150-2 [indeterminate].
Non “Nutica vitellus (Linnaeus, 1758)”. Dillwyn, 1817: 979; Lamarck, 1822:
200; Deshayes, 1832: 601-2; Philippi, 1849: 12-13, pl. 1 , figs 10,
11; Reeve, 1855: pl. 10, figs 39a, 39b; Sowerby, 1883: 9?3?3+, pl. 457,
fig. 41. [Is Nutica stellata Hedley, 19131.
Non “Natica rufa (Born, 1778)”. Brown, 1827: pl. 43, figs 3, 6; Fleming, 1828:
3 19 [indeterminate].
Non “Nuticina rufa (Born, 1778)”. Forbes, 1836: 193 [indeterminate].
Non “Natica (Natica) vitellus (Linnaeus, 1758)”. Tryon, 1886: 29, pl. 8, fig. 60
[Is Natica stellata Hedley, 19131.
Description
Shell globular (shell axis to 55 mm; diameter to 53 mm), unornamented
except for fine subsutural striae; sutures adpressed. Colour light brown, often
white abapically. Diagnostic white (or pale yellow) spiral band running along
.A R K..\BA’T
12
centre of last whorl; subsutural region also may be whitish. Protoconch reddishpurple. Shell interior M hite; outside colour pattern may be seen within. Broad,
semicircular, slightly prosocline aperture; outer lip thin; inner jcolumellar) lip
thickened abapicall) . Umbilicus slightly covered adapically by white parietal
callus, hemiomphalous; umbilicus extending nearly to apex. Umbilical callus
barel! developed; funiculus vestigial. ‘CVhite paucispiral calcareous operculum
w i t h two marginal ribs (the inner broader than the outer) (Fig. 2D). Reduced
third rib sometimes extends partially from the opercular nucleus. Operculum
cdumellar edge distinct11 serrated. Nucleus slightly elevated and may he
encrusted with fine sand grains. Inner curface of the operculum covered with a
thin corneous la) er.
evamzned
‘l’he material in the LSL consists of one box, No. 563, with two shells, the
smaller here selected as the lectotype of,\erzla mtellur Linnaeus, 1758 (Fig. 2A,B),
and the larger referable to L2atzcu ctellnh Hedle), 1913.
This specie\ ma4 not included in the L1.L.L. (Linnaeus, 1764); hence the
material labelled “z~i1ellus” in the Z M V C [two shells in an un-numbered box,
both referable to .\hlicn stellala Hedley, 1913; fig. 2E) were probably subsequently
added to the collection (Odhner 1953: 26). Since Hedley (1913) was merely the
first t o 1 alidatr “ ~ l e l / u / z Ii ~f a r t ) n” 1786) [a non-hinominal work], the original
t>iie material of jtellatu may \\ell be lost.
.\inlei 101
Remurks
The single illustration cited by Linnaeus i1758, 1767j, that of Rumphius
1,1705, 111. 22, fig. Dj, clearly represents the Indo-Pacific .+’atica uilellus (Linnaeus,
1 i.58;, as redescrihed above. However, the figures cited by Born (1778, 1780)
and Gmelin ( 1791 fbr ‘*<:itellus“are of a different species. Born and Gmelin
referred to a shell with a pattern of white spots which is now known as .,Vatica
.sfellnta Hedley. 1913. T h e shell of .2.stellatus is orange with two to three spiral
rows of irregular white ‘oblong to triangular) spots (Fig. 2E); the parietal callus
is purplish-pink and broader than that of .\.. ci/ellus.
(hgraphzcal range
’This species occurs in the central and western portions of the tropical
Tndo-Pacific: China (Shanghai) and Hong Kong, Japan (Tosa) and Okinawa,
Philippines, hfoluccas, Fiji. south\\ ards to Australia (Southport, Queensland to
Exmouth, Il’estern ‘Australia, west to the Ba) of Bengal, the Persian Gulf, and
south to Mauritiui.
~
Genus .\k/icarius DumCril, 1806
111 the <oo/ogze AlnaCbtzgu~, 18061, Dumkril attempted to provide a natural
clddication of the animal kingdom, togethcr with redescriptions of various
t‘ix‘i. Notabl), he distinguished bet\been the hard parts (e.g. skeleton, shell) and
their soft part5. He used the generic name for the shell, and a modified version
for the ‘soft parts’. Thus, in his discussion of .Vulzca Scopoli, 1 7 7 7 (which he
,ittributed to Lamarck I, Dumkril (1806: 164-5) created the term Xatzcarzus for
the animal and reserbed ut’n/zca for the shell. Dumkril did not indicate any species
LINNAEUS’ NATICIDAE
Figure 2. A, Natzca vztellus (Linnaeus, 1758). Lectotype, LSL (No. 563), apertural view. Shell axis
22.2 mm; diameter 22.1 mm. B, Natica vitellus (Linnaeus, 1758). Lectotype, LSL (No. 563), dorsal
view. C, Nerita rufa Born, 1778. Holotype, Naturhistorisches Museum Wien (No. N.1.9), apertural
view. Shell axis 26.5 mm; diameter 26.0 mm. Photograph courtesy Erhard Wawra. D, N Q ~ ~uitellus
CQ
(Linnaeus, 1758), operculum. MCZ 262368 (Ang Hin, Cholburi, Thailand). Operculum 18.6 mm
by 10.4 mm (shell axis 28.9 mm, diameter 27.6 mm). E, N&Q stellala Hedley, 1913. ZMUU
(“uitellus”), apertural view. Shell axis 42.4 mm; diameter 40.2 mm. F, NQtiCariU$ canrena (Linnaeus,
1758). Lectotype, ZMUU (No. 383), apertural view. Shell axis 39.8 mm, diameter 38.6 mm. G ,
Nalicarius canrena (Linnaeus, 1758). Paralectotype, ZMUU (No. 383), dorsal view. Shell axis
35.6 mm, diameter 33.5 mm. H, Naticarius canrena (Linnaeus, 1758), operculum. MCZ 106095
(Boynton, Lake Worth, Florida). Operculum 20.7 mm by 13.1 mm (shell axis 30.6 mm, diameter
28.6 mm).
13
14
A. R. K.iB.47
for either ‘genus’; however, the translation of Froriep (1806: 165) included only
one species for Naticarius, viz. .Nerila canrena Linnaeus, 1758. Later authors have
accepted this as the type species, by subsequent designation (or as the first
included species) (Iredale, 19 16). Despite DumCril’s original intention that
Natica and A“aticarius were two aspects of the same organism, Naticarius has been
regarded as a separate genus from (or subgenus within) Nutica (Woodring,
1957: 85; Majima, 1989: 76).
,Vaticarius canrena (Linnaeus, 1758)
(Fig. 2F-H)
Synonymj
.Nerita canrena Linnaeus, 1758: 776, No. 623 [Figures cited: Rumphius (1 705),
pl. 22, fig. C; Gualtieri ( 1742), pl. 67, figs E, Q R , S, V, X; Argenville ( 1742),
pl. 10, fig. C; Regenfuss ( 1 758), 10, pl. 3, fig. 34. Locality: “0.Asiae”.].
“Verzta canrena Linnaeus, 1758. Linnaeus, 1764: 674, No. 383 [references as in
1758, except that Regenfuss ( 1758) was not included and only the first four
figures of Gualtieri
(1742) were cited. Four varieties were
established.]; Linnaeus, 1767: 1251, No. 715 [Two additional references
cited: Buonanni (1684), figs 224, 228; and Adanson (1757), 1, pl. 13, fig. 3.
Locality given as “0.Asiae, ilfricae”.]; Born, 1778: 410-412; Born,
1780: 396-7, pl. 17, figs. 1, 2; Gmelin, 1791: 3669-70; Turton, 1802:
544-5; Dillwyn, 1817: 975-8; Hanley, 1855: 392-3, 541 [Hanley noted
that six species “at the least” were combined by Linnaeus].
.2iitica ranrena (Linnaeus, 1758). Lamarck, 1816: 11, pl. 453, figs la, lb;
Lamarck, 1822: 199; Deshayes, 1832: 600; Deshayes, 1838: 633-635;
Philippi, 1849: 8-9, pl. 1, figs 5, 6 [“Westindien, bis Rio Janeiro”. Philippi
was apparently the first to recognize that this was solely a Western Atlantic
species.]; Reeve, 1855, pl. 4, figs 14a, 14b; Sowerby, 1883: 79, pl. 455,
fig. 24, pl. 462, fig. 169.
.Vatzca (,Vahca) canrena I Linnaeus, 1758). Tryon, 1886: 20-21, pl. 4, fig. 58.
vliatica (h’alicarzus) canrena (Linnaeus, 1758). Abbott, 1974: 159, pl. 4, fig. 1715.
.Van “.Verita cancrena” [sic] Linnaeus, 1758. Salk Marschlins, 1793: 378; Salis
Marschlins, 1795: 472-3 [Referred to Chemnitz (1781, 5, pl. 186, figs 1860-I),
although the specimens were obtained at Naples. Probably misidentified.]
,Yon “,%atica canrena (Linnaeus, 1758)”. BOSC,1801: 286-7, pl. 28, figs 5-6;
= ,Vatzca ~ t e r c u ~ r n u ~ c a r(Gmelin,
um
1791)].
Van “,Verita canrena Linnaeus, 1758”. Maton & Rackett, 1804: 223-4
[indeterminate].
.Van “.Vatzca ranrenn (Linnaeus, 17581”. hlontagu, 1808: 148-9; Brown, 1827:
pl. 43, figs 13, 16 [?= -4fatica intricata Donovan].
DPJcripLion
Shell globular, (shell axis to 65 mm, shell diameter to 60 mm), unornamented
except for distinct subsutural striae. Spire slightly elevated, sutures distinct,
adpressed; subsutural ridge (“shelf”) pronounced. Protoconch and early
teleoconch yellowish-purple. Considerable polymorphism of the teleoconch
colour patterns, typically a yellow, red or light brown field with four narrow
white spiral bands extending across the last whorl. Abapical and subsutural
LINNAEUS’ NATICIDAE
15
regions may be lighter than the interband regions. Concentric, wavy reddish
brown lines extend across the bands and may form distinctive prosocyrt chevron
marks within the white bands. Spacing and width of the transverse lines can vary
greatly. Shell interior white; outside colour pattern may penetrate partially.
Broad, semicircular, slightly prosocline aperture; outer lip thin; inner
(columellar) lip thickened abapically. Umbilicus nearly filled with broad
umbilical callus (hemiomphalous) ; well-developed funiculus. Distinct notch
between umbilical and parietal calluses, resulting in a sickle-shaped umbilical
groove; inductura glossy white. White paucispiral calcareous operculum with
eight to ten marginal ribs (the outermost thinnest); the columellar edge finely
serrated (Fig. 2H). The nucleus is depressed, somewhat darkened. Inner surface
covered with smooth corneous layer.
Material examined
There is no material in the LSL of this species (Dance, 1967: 21; contra
Jackson, 1913: 43). The material in the ZMUU consists of one tray, No. 383, with
four shells. As Odhner (1953: 19) noted, only two actually represent Natica
canrena. There is also one specimen of the Mediterranean Natica stercusmuscarum
(Gmelin, 1791) [ = “N. millepunctata Lamarck”, of Odhner, 19531 and one
specimen of the Indo-Pacific Natica undulata (Roding, 1798) [ = “ N .zebra
Lamarck”, of Odhner, 19531. This mix-up, however, is not unexpected since the
original descriptions of Jv. canrena were sufficiently broad as to encompass these
latter species [the third and fourth “varieties” of Linnaeus, 1764, correspond to
these taxa]. The larger shell in the ZMUU, tray No. 383, is herein selected as the
lectotype of Nerita canrena Linnaeus, 1758 (Fig. 2F); the smaller shell the
paralectotype (Fig. 2G).
Remarks
Of the numerous figures cited by Linnaeus (1758, 1767), only that of Gualtieri
(1742, pl. 67, fig. V) is clearly referable to Linnaeus’ diagnosis and specimens of
the Western Atlantic Naticarius canrena (Linnaeus, 1758). The other figures of
Gualtieri (1742, pl. 67, figs Q, R, S) and Buonanni (1684) are the Eastern
Atlantic Natica stercusmuscarum (Gmelin, 1791); Adanson ( 1757) is Natica fane1
(Roding, 1798) (Fischer-Piette, 1942: 276-7, pl. 10, fig. 6a-b); and Gualtieri (1742,
pl. 67, fig. X) may represent the Caribbean Natica (Glyphepithema) cayennensis
(Rkcluz, 1850). The figures of Rumphius ( 1705), Argenville ( 1742), Regenfuss
(1758) and Gualtieri (1742, pl. 67, fig. E) do not resemble any one naticid
species [Martens (1902: 114) claimed that Rumphius, pl. 22, fig. C was Natica
chinensis Lamarck = Natica onca (Roding, 1798)].
Geographical range
This common species is found in the tropical Western Atlantic, from North
Carolina (offshore) and Bermuda, southwards to Florida and the Bahamas,
throughout the Gulf of Mexico and Caribbean Sea, Brazil, and possibly south to
Uruguay [MCZ; Kabat, unpublished]. As well, it has an extensive fossil record
in the West Indies and surrounding continental regions [Upper
Miocene-Recent] . A comprehensive synonymy based on fossil records was
compiled by Weisbord (1962: 244-6). Earlier, Woodring had noted the
proliferation of citations of fossil “canrenu” from the Caribbean and commented
16
A.
R. KABBAT
that “it would take an exhaustive study to attempt to find out just what these
fossils represent’’ (b’oodring, 1928: 381). Jung (1964) analysed this species,
along with its fossil subspecies and congeners. Kabat (in preparation) has a full
synonymy of which the foregoing is only a small part.
Genus Polinices Montfort, 1810
’The comprehensive monograph of Denys de Montfort (1810) on the classification of shelled molluscs provided diagnoses for numerous genera, some new,
including the naticids Naticus (p. 218) and Polinices (p. 223). T h e former, with its
type species (original designation) being Nerita canrena Linnaeus, 1758, is a junior
objective synonym of Naticarius Dumtril, 1806. The latter, with type species
(original designation) Polinices albus Montfort, 1810, is the type genus of the
naticid subfamily Polinicinae. The description of Polinices provided by Montfort
is as follows: “Coquille libre, univalve, a spire rtgulikre, relevte, mamille‘e ou
mamelonnke; ayant un ombilic; bouche arrondie, oblongue, tvasie, entikre;
IPvres tranchantes”. Unfortunately, Montfort’s species was poorly illustrated and
the type material has not survived. However, it has been regarded as a junior
synonym of Jerita mammilla Linnaeus, 1758 (cf. Majima, 1989: 42-3) and is
herein treated as such.
Polinices mammilla (Linnaeus, 1758)
(Fig. 3A, B)
Synonymy
,Verita mammilla Linnaeus, 1758: 776-7, No. 627 [Figures cited Colonna (l606),
pi. 52, fig. ult; Lister (1685-92), 4, f. 5, c. 3, t. 1, f. 3, 4; Rumphius (1705),
pl. 22, fig. “E” (corrected to “F” in Linnaeus ( 1 764, 1767)); Gualtieri (1742),
pl. 67, figs C, D; Argenville (1742), pl. 10, fig. X. Locality: “Barbados”.].
h‘erita mammilla Linnaeus, 1‘758. Linnaeus, 1764: 675, No. 386 [references to
Colonna (1606) and Lister (1685-92) omitted.]; Linnaeus, 1767: 1252,
No. 719 [added reference to Seba (1758), 3, pl. 38, figs. 9, 10. Locality given
as “Barbados, Xlexandriae”.]; Born,
1778: 415-6; Born,
1780:
399-400; Gmelin, 1791: 3672; Turton, 1802: 546; Dillwyn, 1817: 984-5;
Hanley, 1855: 396-7, 542 [noted that several species were combined and that
this “appears to be intended for the common imperforated snow-white
Oriental shell . . .”].
Albula mammilla (Linnaeus, 1758). Roding, 1798: 20.
*+’&a
mamilla [sic] (Linnaeus, 1758). BOSC,1801: 288; Lamarck, 1822: 197
[“l’Ocean des grandes Indes”]; Gray, 1826: 482; Deshayes, 1832:
599; Deshayes, 1838: 630-1; Philippi, 1852: 31-32, pl. 4, figs 7, 8
[“Ostindien”]; Reeve, 1855: pl. 7, figs 27a, 27b [“Island of Luzon,
Philippines. . .This we take to be the original type of the old Linnaean Nerita
mamilla”.]; Sowerby, 1883: 85, pl. 456, figs 29, 30.
,Vatica mammilla (Linnaeus, 1758). Lamarck, 1816: 1 1, pl. 453, figs 5a, 5b.
Mamillaria tumidus Swainson, 1840: 345 [Referred to Chemnitz (1781, 5, pl. 189,
figs 1928-1931). Lectotype, Zoologisk Museum, Kobenhavn (selected and
figured by Cernohorsky, 1974: 172, fig. 44)].
LINNAEUS’ NATICIDAE
17
Natica pyrzformis Rtcluz, 1844b: 21 1 [Not illustrated; locality given as “110-110,
island of Panay. . .Huan River, Australia”.] Philippi, 1852: 60, pl. 9, fig. 8;
-Reeve, 1855: pl. 5, fig. 16.
Natica albula Rtcluz, 1851: 194-197 [Cited from Rumphius (1705, pl. 22, fig. F);
locality given as “les c6tes d’Ele Wallis, Amboine, etc.”].
Naticaponderosa Philippi, 1852: 32, pl. 4, figs 9, 10 [Referred to Chemnitz (1781,
5, pl. 189, figs. 1930-1931); locality not known].
Natica cygnea Philippi, 1852: 80-1, pl. 12, fig. 6 [Locality not stated].
Natica (Mamma) mamilla [sic] (Linnaeus, 1758). Tryon, 1886: 49, pl. 16,
figs 46, 48.
Polinices (Polinices) tumidus (Swainson, 1840). Cernohorsky, 1971: 191-193,
figs 45, 47-50; Cernohorsky, 1974: 172, fig. 44; Kilburn, 1976: 856-857,
fig. 15.
Non “Nerita mammilla Linnaeus, 1758”. Salis Marschlins, 1793: 380; Salis
Marschlins, 1795: 474 [Referred to Chemnitz (1781, 5, pl. 189, figs 19281931). Probably a misidentification since the specimens were obtained at
Naples where no species resembling mammilla occurs].
Non “Natica mammilla (Linnaeus, 1758)”. Brown, 1827, pl. 43, figs 9, 11 (not
naticid).
Description
Shell pyriform (shell axis to 57 mm, diameter to 44 mm), unornamented,
moderately high spired, with distinct flush sutures. Glossy white under dark
reddish-brown periostracum. Protoconch reddish to black. Aperture slightly
prosocline, semicircular; peristome simple, outer lip thin, inner (columellar) lip
slightly thickened basally. Shell interior glossy white. Distinctive umbilical callus
completely (or nearly so) filling umbilicus, cryptomphalous; only a vestigial
umbilical groove abapically (in some specimens). Parietal callus fused with
umbilical callus. Line of umbilicus runs continuously from abapical columellar
lip to adapical edge of the outer whorl. Corneous paucispiral pale-brown
opercul u m.
Material examined
There is no material in the LSL referable to Polinices mammilla (Dance, 1967:
22); but there are two trays in the ZMUU, both numbered No. 386, each
containing one shell. The smaller yellow shell is a specimen of Polinices aurantius
(Roding, 1798), as noted by Odhner (1953: 19). The larger white shell is
referable to the Indo-Pacific Polinices marnmilla (Linnaeus, 1 758) (Odhner,
1953: 19), as herein restricted and is here selected as the lectotype of Nerita
mammilla Linnaeus, 1758 (Fig. 3A,B). In his M.L.U. description, Linnaeus
( 1764) included the phrase “lactea aut lutea”, which could encompass both
specimens in the ZMUU. As noted in the historical background of the genus
Polinices, the type species Polinices albus Montfort, 1810 has been regarded as a
junior synonym of mammilla, despite the sketchy description and lack of type
material for albus (not found in the Mustum National d’Histoire Naturelle,
Paris). In order to achieve nomenclatural stability, I designate the lectotype for
mammilla as the neotype for albus, thus making albus an objective junior synonym
of mammilla. If this is not done, some authors might reject albus as a nomen
18
A.
K. KABAT
dubium and thereby discard entirely the genus Polinices which is probably the
second most used naticid generic name (after .iVatica).
Remarks
Linnaeus’ original locality data, “Barbados” (Linnaeus, 1758) and
“Barbados, Alexandriae” (Linnaeus, 1767) does not correspond with any
Atlantic species and has misled some authors (e.g., Marincovich, 1977: 246) into
concluding that mammilia could not be an Indo-Pacific species.
Most of the illustrations cited by Linnaeus represent the Indo-Pacific Polinices
mammilla; the better figures are those of Colonna (1606, pl. 52, fig. ult); Lister
(1685-92, pl. 571, fig. 22); Gualtieri (1742: pl. 67, fig. C) and Seba (1758, pl. 38,
figs 9, 10). The other illustrations are indeterminate, and the second figure of
Cualtieri (1742, pl. 67, fig. D) appears to be of the naticid genus Mummilla
Schumacher.
’This taxon, in addition to being frequently misspelled mami mil la^'), has been
rejected by some recent authors in favour of Mamillaria tumidus Swainson, 1840.
Cimmhorsky (1971: 191; 193) stated that “. . .,Verita mamilla [sic]. . .may be an
earlier riame for Polinices lacteus (Guilding) from the West Indies” and that
“‘Taxonomic stability would be best served by discarding Nerita mamilla [sic] as a
nomen dubium”. The cordusion is illogical: mammilla has been used repeatedly
ovcr the years whereas tumidus was rarely (if ever) used between its description
and 1971! Analysis of the original description, cited figures, and type material
leads t o the conclusion that mammilla is, in fact, a senior synonym of tumidus arid
it is herein restricted to the white-shelled, closed-umbilicus Indo-Pacific Polinices
(“lactea” of Linnaeus, 1758) while the eponymous aurantius Roding, 1798 is used
for the Indo-Pacific Poiinices with golden-orange shells (“lutea” of Linnaeus,
1758). The shells of the closely related PolinicesJIemingiana (Rtcluz, 1844) can be
separated from mammiila by the umbilical groove, broader aperture, and more
oblique outline in the former (Cernohorsky, 1971: figs 45, 46).
Geo<graphicalrange
’This species is one ctf the most common naticids throughout the tropical
Indo-Pacific. It is known from the Hawaiian Islands, south-west through
Polynesia, Melanesia, Micronesia, Malaysia, Philippines, north to Japan
(Oshima, Bashio), New Guinea, Norfolk and Lord Howe Islands, Australia
[Brisbane, Queensland to Geraldton, Western Australia), westwards to the Bay
of Bengal, Ceylon, and Arabian Sea, Diego Garcia, Mauritius, Madagascar and
from the Persian Gulf to Durban.
Genus ,Veverita Risso, 1826
Kiss0 (1826), in his monograph on the natural history of Europe (particularly
the Mediterranean Sea), described two naticid genera, Nacca and Neverita. The
former is a junior subjective synonym of Natica Scopoli, 1777; the latter is
currently placed in the naticid subfamily Polinicinae, either as a full genus
(herein] or as a subgenus of Polinices Montfort, 1810. The type species, by
monotypy, is Neverita josephinia Risso, 1826 from the Mediterranean. Risso’s
diagnosis of this genus was:
LINNAEUS’ NATICIDAE
19
Figure 3. A, Polinices mammilla (Linnaeus, 1758). Lectotype, ZMUU (No. 386), apertural view. Shell
axis 43.5 mm, diameter 35.0 mm. B, Polinices mammilla (Linnaeus, 1758). Lectotype, ZMUU
(No. 386), dorsal view. C, NeNeverita albumen (Linnaeus, 1758). Lectotype, ZMUU (No. 385),
apertural view. Shell axis 34.0 mm, diameter 42.0 mm. D, Neuerita albumen (Linnaeus, 1758). MCZ
243960 (Tayabas, Luzon, Philippines), dorsal view. Shell axis 24.4 mm, diameter 33.2 mm.
“Testa ovulata; anfractubus depressis, nullomodo elevatis; sutura indistinctissima, peritrema postice ad sinistram valde incrassatum, proeminens,
umbilicum claudens; operculum cartiloginosum” (Risso, 1826: 149).
Xeuerita albumen (Linnaeus, 1758)
(Fig. 3C,D)
Synonymy
Nerita albumen Linnaeus, 1758: 776, No. 626 [Figures cited: Lister (1685-92), f. 5,
c. 3, f. 1, 2; Rumphius (1705), pl. 22, fig. B; Gualtieri (1742), pl. 67, figs. A, B;
Argenville (1742), pl. 10, fig. T; Regenfuss (1758), 20, pl. 5, fig. 54. Locality:
“0.Asiae”.]
20
4 . R. KABAT
Aerita albumen Linnaeus, 1758. Linnaeus, 1764: 675, No. 385 [references to
Lister (1685-92) and Regenfuss (1 758) omitted]; Linnaeus, 1767: 1252,
No. 718; Born, 1778: 415; Born, 1780: 399; Gmelin, 1791: 3671-2;
’Furton, 1802: 545-6; Dillwyn, 1817: 984; Hanley, 1855: 395-6, 542 [suggested that Argenville (1742) should be pl. 10, fig. V (rather than fig. T) and
Regenfuss ( 1758) should be pl. 3, fig. 34 (rather than fig. 54)].
,Vatica albumen (Linnaeus, 1 758). BOSC, 1801: 288; Lamarck, 1822:
196 7; Deshayes, 1832: 598; Deshayes, 1838: 627--8; Philippi, 1852: 2930, pl. 4, figs 3--4; Reeve, 1855: pl. 8, figs 31a, 31b; Sowerby, 1883: 78,
pl. 458, fig. 57.
,Valica ( M a m m a ) albumen (Linnaeus, 1758). Tryon, 1886: 47, pl. 20, fig. 5.
Polinices (Neverila) albumen (Linnaeus, 1758). Cernohorsky, 1971: 195-196,
figs57, 58; Kilburn, 1976: 859.
Descriplion
Shell broad (shell axis to 34 mm, diameter to 42 mm), low-spired, smooth and
glossy. Spire depressed, adapical aspect nearly flat. Sutures adpressed, almost
flush. Rich chestnut brown colour; protoconch and early teleoconch off-white as
is a narrow subsutural region. Indistinct whitish periostracum may be present
(Fig. 3D). Shell interior white, except at the outer edge where the brown shows
through. Exceedingly prosocline semicircular aperture, outer lip thin, inner lip
thickened. Umbilicus deep, nearly cryptomphalous, largely filled with the glossy
white callus which lacks a transverse groove; shallow notch between umbilical
and parietal inductura. Funicle may be irregularly twisted as it ascends the
umbilicus. Adaxial rim of umbilicus flattened, white umbilical groove broad,
semicircular. Corneous paucispiral dark brown operculum.
Material examined
There is no material in the LSL (Dance, 1967: 21; contra Jackson, 1913: 43).
It is not clear what has happened to the specimens referred to by Hanley (1855:
395). The material in the ZMUU consists of a one tray, No. 385, with two
conspecific shells of Nererila albumen (Linnaeus, 1 758); the larger herein selected
as the lectotype (Fig. 3C) and the smaller the paralectotype.
Remarks
As Hanley (1855) noted, the illustrations cited by Linnaeus ( 1 758, 1767) are
not of great assistance in determining this species. None of the illustrations agree
with Linnaeus’ type material or original description; although those of
Rumphius ( 1705) and Gualtieri ( 1 742) are probably of Neverila species. Kilburn
(1976: 859) stated that Hanley (1855: 395) had ‘designated’ the type figure as
Rumphius ( 1 705, pl. 22, fig. B); however, Hanley had merely suggested that
“the majority of conchologists have preferred to retain the appellation for the
shell delineated by Rumphius.. .” without formally designating it as the type for
h: albumen.
Geographical range
This uncommon species is found in the central and western portions of the
tropical Indo-Pacific: Japan (Kagoshima Bay), Okinawa, Philippines, Malaya,
LINNAEUS’ NATICIDAE
21
Moluccas, Solomons, Fiji, New Caledonia, Australia (Lindeman Island,
Queensland to Dampier Archipelago, Western Australia), westwards to
Mauritius, Madagascar, and South Africa (Natal).
Nerita glaucina Linnaeus, 1758 (Nomen dubium)
Nerita glaucina Linnaeus, 1758: 776, No. 624 [Figures cited: Lister (1685-92), f. 5,
c. 1, t. 5, f. 1; Rumphius (1 705), pl. 22, fig. A; Gualtieri (1 742), pl. 67, figs. M,
P, T. Locality: “0.Europaeo”.].
Nerita glaucina Linnaeus, 1758. Linnaeus, 1761: 533, No. 2 197 [the only cited
illustration is Lister (1678), 163, pl. 3, fig. 10.1; Linnaeus, 1764: 674, No. 384
[reference to Lister (1685-92) omitted.]; Linnaeus, 1767: 1251-2, No. 716
[references to Gualtieri (1 742) and Rumphius ( 1705) unchanged; Lister
(1678) p. 163, pl. 3, fig. 10; Adanson (1757), 1, pl. 13, fig. “14” (error for 4)];
Born, 1778: 412-3; Born, 1780; 397, pl. 13, figs. 20, 21; Gmelin, 1791;
3671; Turton, 1802: 545; Hanley, 1855: 393-4, 541; pl. 3, fig. 5 [Hanley
noted that several unrelated naticid species were confused under this name,
concluded that this species is from Gibraltar and Algeria, and suggested that it
may be close to Natica collaria Lamarck, 18221.
Natica glaucina (Linnaeus, 1758). Lamarck, 1822: 1 96; Deshayes, 1832:
597-8; Deshayes, 1838: 625-7 [Deshayes referred the Swedish records to
Natica monilifera Lamarck, 1822; and the Mediterranean to Natica olla Serres,
1829 ( = Neverita josephinia R h o , 1826).]
Non “Nerita glaucina Linnaeus, 1758”. Pennant, 1777: 140, pl. 87, fig. 141;
Dillwyn, 1817: 978-9 [ = Euspira catena (Costa, 1778)].
Non “Nerita glaucina Linnaeus, 1758”. Costa, 1778: 83-84, pl. 5,
fig. 7; Donovan, 1799: pl. 20, figs 1-2; Montagu, 1803: 469-470; Maton
and Rackett, 1804: 224-5; Fleming, 1828: 319 [ = Euspira alderi (Forbes,
183811.
Non “Nerita glaucina Linnaeus, 1758”. Salis Marschlins, 1793: 378; Salis
Marschlins, 1795: 473; Reeve, 1855: pi. 3, fig. 8 [“Mediterranean”]. [Is
Neverita josephinia Risso, 1826.1
Non “Natica glaucina (Linnaeus, 1758)”. Philippi, 1845: 44-45, pl. 2, figs. 10, 11
[“Mare Germanicum et M. Mediterraneum”]; Philippi, 1852: 100-101,
pl. 14, figs 7, 8 [“die Kusten Europas von Scandinavien bis Neapel”]. Is
Euspira alderi (Forbes, 1838).
,I
Material examined
The material in the LSL consists of one box, No. 562, with six shells. These six
specimens are a mixed assortment of several European species, including Natica
stercusmuscarum (Gmelin, 179 1 ), Euspira alderi (Forbes, 1838) and E. catena (Costa,
1778).
The material in the ZMUU consists of one tray, No. 384, with two conspecific
shells, both referable to Neverita. Odhner (1953: 19) suggested that they represented the Indo-Pacific Natica ampla Philippi, 1849 [ = Neverita didyma (Roding,
1798)]. The conclusion herein is that neither lot represents valid typological
material for this Linnaean name: they do not agree with the original description
or the cited figures.
22
A R KABr\T
Remarks
Cnfortunately, the early illustrations cited for this species are singularly
uninformative: those of Lister (1685-92) and Rumphius ( 1 705) are indeterminate ise\.eral authors used “Rumphius, pl. 22, fig. A” for X . vitellus instead of
“fig. D”) while those of Gualtieri (1742) are probably referable to Natica;
Adanson (1757, pl. 13, fig. 4) is of the West African Natica fulminea (Gmelin,
1791) (Fischer-Piette, 1942: 278-9). The conclusion is that none of the initially
cited illustrations adequately match the original descriptions or the type
material. Nor did Tryon (1886: 33) solve the problem: he stated that the
“glaucina” from the Mediterranean was Neverita josephznza Risso, 1826; and also
{ 1886: 41 I that “glauczna” in part [i.e. from Northern Europe] was Natica alderi
Forl>es, 1838.
The inescapable conclusion is that ”Verita glaucina is a nomen dubium, as the
descriptions, cited illustrations, and type material do not agree on any one
naticid species ( o r even one naticid genus!). I t appears that Linnaeus had, at
various times, confounded shells of Euspira from Northern Europe and
indeterminate Mediterranean or Indo-Pacific shells of Neverita. Although
Odhner 11953) considered ‘glauczna’ to be a Neverila, the species, cannot be
definitively treated as a senior synonym of either the Mediterranean Neverita
psephznza Risso, 1826 or the Indo-Pacific ,Veverita dzdyma (Roding, 1798). Both of
these latter names h a w been frequently used over the last 150 years, and they
represent clearly-defined, common and M ell-known species. It seems best, therefixc, to retain their usage and to reject Linnaeus’ Nerita glaucina as a nomen
dubzum. This will have little effect on naticid taxonomy as glauczna has been used
rare11 fif at all) in this century.
ACKSO\\’LEDGEMEXI‘S
Solene Morris [British Museum (Natural History)] greatly assisted my visit to
the LSL in June, 1987, and provided much helpful discussion with respect to the
Linnaean shell collections and their provenance. Dr Anders Warkn and Prof.
Wke Franztn [Naturhistoriska Riksmuseet, Stockholm] arranged my visit to the
ZMUU in July, 1987. At the ZMC‘U, Dr Sten Jonsson kindly assisted my
examination of the specimens of the Museum Ludovicae Ulricae, as well as those
in the general collections. Dr Yves Finet [Muskum d’Histoire Naturelle de
GenPve] allowed me to examine the naticid type specimens of Lamarck and
RCcluz. Dr Erhard Wawra [Naturhistorisches Museum Wien] provided photographs of the type material of ,Verzta rufa Born, 1778. For data on the geographical ranges, I am grateful to the following curators for providing access to the
museum collections: I . Loch and CY. F. Ponder (AMS), R. Kawamoto (BPBM),
K . J . Boss (MCZ) and S. Slack-Smith (CYAM). T h e following individuals have
critically reviewed this manuscript and their comments have been most helpful:
K. J. Boss, A. J. Kohn, R. E. Petit, and an anonymous reviewer.
LINNAEUS' NATICIDAE
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