Type specimens of Conus (Mollusca: Gastropoda) in the Zoological

type specimens of conus
97
Type specimens of Conus (Mollusca: Gastropoda) in the Zoological
Museum of the University of Copenhagen: A Historical Account
ALAN J. KOHN
Steenstrupia
Kohn, A. J. Type specimens of Conus (Mollusca: Gastropoda) in the Zoological Museum of the University of Copenhagen: A Historical Account. – Steenstrupia 30 (2): 97–113. Copenhagen, Denmark.
April 2009. ISSN 0375-2909.
This paper reviews the sources and history of the 20 type specimens of the large marine gastropod
genus Conus in the Zoological Museum of the University of Copenhagen (ZMUC), the authors and
their publications, and the present status of the nominal species. Sixteen are primary or name-bearing
type specimens that served as the bases for 18 species of Conus; one of these is also a syntype of a 19th
described species. The others are three paratypes and one paralectotype. Most derive from the collections of Counts Adam and Joachim Moltke and Lorenz Spengler, and they have resided in the Museum
since the Museum Moltkianum came to the University nearly 200 years ago. Their publications date
back even farther, to Chemnitz’s description of the famous Conus gloriamaris in 1777. Eleven of the
species were described in the 18th Century, nine in the first half of the 19th, and two, C. knudseni and
C. sorenseni, in 1982. Today only five of the nominal species-group names are considered valid, the
others having been sunk in synonymy.
Keywords: Conus, type specimens, Zoological Museum, Copenhagen
Alan J. Kohn: Department of Biology, University of Washington, Seattle, Washington 98195, U.S.A.
E-mail: [email protected]
INTRODUCTION
The first major natural history museum in Copen­
hagen was that of Ole Worm (1588–1654), and
a few objects from it remain to this day (Wolff
1999). King Frederik III (1609–1670) was an inspired devotee who developed his own col­lection
and after Worm’s death acquired the col­lec­tion
for his own museum in Christiansborg Castle.
However, this made the collections less accessible to others, and University naturalists began
to build up their own collections in the late 17th
and early 18th centuries. Unfortunately, these collections of animal specimens were destroyed by
fire in 1728 (Wolff 1999).
One year after the tenth edition of Linnaeus’s
Systema Naturae (1758) appeared, Count Adam
Gottlob Moltke (1710–1792), established the
Natural- og Husholdnings-Cabinettet (The Nat­
uralia and Housekeeping Cabinet), a college at
Charlottenlund. He donated some of his own
extensive collections from the Danish tropical
colonies, and also acquired Peter Forsskål’s
Steenstrupia 30 (2): 93–113.
Arabian collections for this museum. Moltke
was also busy in other pursuits, as he is said to
have fathered 22 sons, including five cabinet
mini­sters, four ambassadors, and two generals.
One son, Count Joachim G. Moltke (1746–1818),
inherited his father’s museum, the Museum
Moltkianum. He later became Prime Minister,
whereupon he presented it to the University, in
1810 (Bruun 1945, Wolff 1999).
Fifteen of the 20 Conus type specimens in the
Zoological Museum, University of Copenhagen
(ZMUC) were illustrated in several volumes of
the Neues Systematisches Conchylien-Cabinet, a
voluminous and authoritative work published be­
tween 1769 and 1795 and begun by the Hamburg
physician Friedrich Heinrich Wilhelm Martini
(1729–1778). Martini published the first three
volumes, in 1769, 1773, and 1777. After his
death, Mrs. Martini and her advisor selected Johann Hieronymus Chemnitz (1730–1800) to continue the series. Chemnitz, a clergyman born in
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a. j. kohn
Magdeburg, had moved to Copenhagen in 1768
as pastor to the German church there, bringing
with him his extensive personal shell collection.
He published Volumes 4–11 between 1780 and
1795. In all, this monumental series comprised
4008 pages and 406 colored plates. Cernohorsky
(1974) gives details of the pub­lication history
of these volumes and the artists and engravers
responsible for their illustration.
Martini and Chemnitz carefully described and
accurately illustrated many new species of mol­
luscs in the Neues Systematisches ConchylienCabinet, including numerous Conus species in
Volumes 2 (Martini 1773), 10 and 11 (Chemnitz
1788, 1795). The figures in this work are suffi­
ciently accurately rendered to be unequivocally
identifiable as the type specimens of most of the
Conus species discussed here. However, the no­
menclature in these volumes was not consist­ently
binominal, and the International Commis­sion on
Zoological Nomenclature placed the entire work
on the Official List of Rejected Works in Zoo­logy
(Hemming 1958: p. 5), rendering all of the species names unavailable.
Martini and Chemnitz both described and
illu­strated shells from their own collections in
their volumes. Both collections were sold at
auction after their owners’ deaths. Other speci­
mens illu­strated in the Neues Systematisches
Conchylien-Cabinet derive from the extensive
collections made by the Hamburg physician
Joachim Frie­drich Bolten (1718–1796) and by
Lorenz Spengler (1720–1807). Spengler, originally from Schaff­hausen, Switzerland, emigrated
to Denmark in 1743 and became cabinet-maker
to the king and a close friend of Chemnitz. Spengler was also an ivory turner at the Danish court
and an expert crafter of artificial teeth. Income
from the latter activity allowed him to acquire
a very extensive shell collection (Cernohorsky
1974). He obtained some specimens from the
Bolten col­lection and probably some from
Chemnitz. Specimens from Spengler’s collection
are marked “Sp” in ink within the aperture (see
Pl. 2, Figs. 25, 28, 35). The Royal Natural History
Museum in Copenhagen purchased Spengler’s
collection in 1805, two years before he died. This
Museum had been established in 1789 by the
Natural History Society of Copenhagen, largely
at the initiative of Peter Christian Abildgaard
(1740–1801). Abild­gaard, a physician, naturalist
and the founder of the Royal Veterinary College, contributed im­portantly to copepodology
and parasitology. He first demonstrated the life
cycle of a cestode parasitic in marine birds with
stick­lebacks as the intermediate host (Buchmann
1997). Abildgaard became the author, editor
and publisher of the faunistic series Zoologica
Da­nica after the death of O.F. Müller in 1784
(An­ker 1950). Abildgaard chaired a committee
to form the Royal Natural History Museum, and
it replaced that of the Natural History Society.
In 1862, this museum was incorporated with the
University Zoological Museum, thus joining the
Moltke collection, and the new museum building
opened in Krystalgade in 1870 (Wolff 1999).
Abildgaard’s nephew, Hans Severin Holten
(1770–1805), became editor of Zoologica Da­
nica after his uncle’s death in 1801. Holten
was also curator of the collection of the Natural
His­tory Society of Copenhagen, prior to its con­
version to the Royal Natural History Museum,
and he tutored Prince Christian Frederik in na­
tural his­tory. Later King Christian VIII (1786–
1848), the prince became committed to natural
history under Holten’s influence, and amassed
an important collection, now also in the ZMUC.
Specimens from King Christian VIII’s collection
are indicated by the designation “CVIII” written
in the aperture of the shells. Holten also prepared
the sale catalogue of the Chemnitz collection, the
Enum­eratio Systematica Conchyliorum Beat J.
H. Chemnitzii (Holten 1802), in which he intro­
duced more than 60 new species-level taxa of
molluscs, including six of Conus (Winckworth
1943, Kohn 1992).
In his earlier historical account of types in
the ZMUC, Cernohorsky (1974) listed types
of only 60 species of Mollusca, including only
two of Conus. We located type specimens of 21
nominal Conus species discussed here during my
1958 and 2007 visits to the ZMUC. Two were
de­scribed after Cernohorsky’s (1974) study. This
report reviews the sources and history of those
specimens and illustrates them. Most were de­
scribed as new during the period just reviewed,
predominantly between 1777 and 1850. In addi­
tion, the previously designated lectotype of one
species cannot be demonstrated to have been a
syntype and should lose its status as lectotype.
type specimens of conus
Following the species accounts, a table sum­
marizes the nominal species, their type speci­
mens, and their currently accepted names.
99
THE TYPE SPECIMENS AND THEIR
IDENTITIES
1992; Filmer 2001) have concluded meets the
criteria of binominal nomenclature.
Chemnitz studied a single specimen, now
regarded as the holotype of C. gloriamaris (Pl.
1, Figs. 1–3). He borrowed it from Count A. G.
Moltke, who had purchased it from a collection
in Holland about 1757. The specimen came to
the University of Copenhagen in 1810 with the
Museum Moltkianum, was later in King Christian VIII’s collection, but after his death in 1848
returned to the University where it has been ever
since (Bruun 1945).
The holotype (Pl. 1, Figs. 2, 3) is 92x35 mm.
A long scar where the animal repaired its shell
after an unsuccessful predation attempt is visible
at the left of Fig. 2. In the original description
Chemnitz (1777: 324–325) described this scar in
detail, clearly indicating that the specimen is the
one Chemnitz described. This is despite the fact
that his original figure, drawn but not engraved
by Franz Michael Regenfuss (1713–1780), the
engraver to the King of Denmark, artistically
“repaired” the shell so as not to show the repair
scar (Fig. 1) (Bruun 1945).
Conus gloriamaris Chemnitz, 1777 is a valid
species, occurring in the western tropical Pacific
Ocean from Indonesia and the Philippines to
Samoa (Röckel, Korn & Kohn 1995).
Conus gloriamaris Chemnitz, 1777
Conus amadis Gmelin, 1791
Material examined:
Holotype of Conus gloriamaris: ZMUC-GAS-114.
Material examined:
1 specimen: ZMUC-GAS-350.
Remarks
Remarks
The noted Danish oceanographer Anton Fr.
Bruun (1945) thoroughly investigated the taxo­
nomic history of Conus gloriamaris and clearly
established Chemnitz as the original author of the
available and valid name of the species. With the
exception of this one, all of the species Chemnitz described were in the Neues Systematisches
Conchylien-Cabinet (1780–1795), so his new
species names are nomenclaturally unavailable
(Hemming 1958). However, Chemnitz (1777)
described C. gloriamaris in a separate pub­li­
cation in a way that most authors who have
investigated the matter (Melvill 1885, 1887;
Tomlin 1937; Jutting 1938; Bruun 1945; Kohn
Gmelin (1791) based Conus amadis on postLinnaean publications showing six illustrations
that he considered to represent this previously
undescribed species. The most accurate of the
figures he cited are those of Chemnitz (1788: pl.
142, figs. 1322, 1323). They show two views of
a specimen from the Nicobar Islands originally
from the collection of Count A. G. Moltke (Pl. 1,
Figs, 4, 5). In 1958, Dr. Jørgen Knudsen located
a specimen of the same size and similar color
pattern (Pl. 1, Figs. 6, 7) in the ZMUC, and
Kohn (1966, 1992) designated it lectotype of C.
amadis Gmelin. However, recent comparison of
this spec­imen with Chemnitz’s figures indicates
METHODS
The accounts of nominal species with type spec­
imens in the ZMUC are arranged chronologi­
cally. Each presents the evidence for considering
the specimen a holotype, lectotype, syntype,
para­type or paralectotype and summarizes the
history of the specimen in the ZMUC. Next,
taxonomic evidence is assessed to determine
whether the described species is valid, a syn­onym
of a previously described species, or an unused
senior synonym that cannot be used because of
provisions of the ICZN. This infor­mation is then
used to conclude the correct present name of each
species. Each specimen is illustrated with digital
images made at the ZMUC in 2007, together with
a reproduction of its ori­g­inal published figure.
Measurements of shells are given as length x
maximum diameter.
Pl. 1, Figs. 1–3
Pl. 1, Figs. 4–7
100
a. j. kohn
that the lectotype is a different specimen from
that illustrated (cf. Figs. 4, 5 and 6, 7). Despite
the nearly identical size, several differences in
shape, particularly at both ends of the aperture,
and color pattern, particularly on the dorsal side,
support this conclusion.
Although I formerly (Kohn 1966, 1992)
con­sidered the specimen shown in Figs. 6, 7
a syn­type of C. amadis and thus eligible to be
de­s­ignated as lectotype, neither the accounts of
Chemnitz (1788) nor Gmelin (1791) document
this. Present evidence is thus not sufficient to
regard the specimen as a syntype. Although it
has not been “demonstrated that a specimen de­
s­ignated as a lectotype is not a syntype” in the
wording of Art. 74.2 of the ICZN (1999), the
absence of positive evidence indicates the spec­
imen should lose its status of lectotype. There­fore
I now designate figs. 1322 and 1323 in Chemnitz
(1788: pl. 142) as representations of the lectotype
of Conus amadis Gmelin, 1791. It is a valid species, occurring throughout the Bay of Bengal
(Röckel, Korn & Kohn 1995).
Remarks
Conus solidus Gmelin, 1791
Remarks
Pl. 1, Figs. 8–10
Material examined:
Holotype of Conus solidus: ZMUC-GAS-351 (= C. mappa
(Lightfoot, 1786)).
Remarks
As Kohn (1992) noted, “the name, cited figure
and diagnosis are all based exclusively on a
single specimen” originally from the Moltke
col­lection and later in that of King Christian VIII
that was described by Chemnitz (1788: p. 59, pl.
141, fig. 1310 (Pl. 1, Fig. 8). This specimen (Pl. 1,
Figs. 9, 10) is the holotype of C. solidus Gmelin,
1791 (Vink & von Cosel 1985, Kohn 1992).
C. solidus is a synonym of C. mappa (Light­
foot, 1786) (Vink & von Cosel 1985, Kohn
1992), most likely of the nominotypical sub­
species, from Trinidad or Tobago (Vink & von
Cosel 1985).
Conus tribunus Gmelin, 1791
Pl. 1, Figs. 11–13
Material examined:
Lectotype of Conus tribunus: ZMUC-GAS-168 (= C. centurio Born, 1778).
Gmelin based Conus tribunus partly on fig. 655
of Pl. 59 in Martini (1773) (Pl. 1, Fig. 11). Kohn
(1966, 1992) designated the shell represented
in that figure, an unmarked specimen from the
Spengler collection present in the ZMUC, as
lectotype of that nominal species (Pl. 1, Figs.
12, 13).
Although evidently not known to Gmelin, this
species had been described thirteen years earlier
as C. centurio by Born (1778), and C. tribunus
Gmelin, 1791 is thus a junior synonym of C.
centurio (Kohn, 1992). C. centurio is a western
Atlantic species, occurring from Jamaica to Bar­
bados and along the South American coast from
Colombia to Rio de Janeiro, Brazil.
Conus cedonulli dominicanus Hwass in Bruguière, 1792
Pl. 1, Figs. 14–16
Material examined:
Lectotype of Conus cedonulli dominicanus: ZMUC-GAS-352
(= C. curassaviensis Vink & von Cosel, 1985).
In Bruguière (1792: 603) Conus cedonulli domi­
Plate 1: 1. Conus gloriamaris Chemnitz. Original figure,
reproduced from Chemnitz (1777: pl. 8, fig. A). – 2, 3. Conus
gloriamaris Chemnitz. Holotype (ZMUC-GAS-114). 92x35
mm. – 4, 5. Conus amadis Gmelin. Representation of lecto­­
type, reproduced from Chemnitz (1788: pl. 142, figs. 1322,
1323). – 6, 7. Conus amadis Gmelin. Specimen formerly
designated lectotype by Kohn (1966, 1992), but lacking documentation that it was a syntype; see text (ZMUC-GAS-350).
78x41 mm. – 8. Conus solidus Gmelin (= C. mappa [Lightfoot, 1786]). Illustration of holotype reproduced from
Chemnitz (1788: pl. 141, fig. 1310). – 9, 10. Conus solidus
Gmelin (= C. mappa mappa [Lightfoot, 1786]). Holotype
(ZMUC-GAS-351). 51x29 mm. – 11. Conus tribunus Gmelin
(= C. centurio Born). Illustration of lectotype reproduced
from Martini (1773: pl. 59, fig. 655). – 12, 13. Conus tribunus
Gmelin (= C. centurio Born). Lectotype (ZMUC-GAS-168).
52x31 mm. – 14. Conus cedonulli dominicanus Hwass in
Bruguière. Illustration of lectotype reproduced from Chemnitz (1788: pl. 141, fig. 1306). – 15, 16. Conus cedonulli
dominicanus Bruguière (= C. curassaviensis Vink & von
Cosel, 1985). Lectotype (ZMUC-GAS-352). 42x21 mm. –
17. Conus cinereus Hwass in Bruguière. Illustration of paralectotype (syntype of Hwass’s Variety C) reproduced from
Chemnitz (1788: pl. 142, fig. 1319). – 18, 19. Conus cinereus
Hwass in Bruguière. Paralectotype (ZMUC-GAS-353, syntype of Hwass’s Variety C). 43x21 mm.
type specimens of conus
101
102
a. j. kohn
nicanus is described as one of nine infraspecific
taxa of C. cedonulli. Linnaeus (1767) originally
described C. cedonulli as an infraspecific form of
C. ammiralis Linnaeus, 1758, but it is now con­
sidered a valid species (Kohn 1992), oc­curring
in the Caribbean region. Because neither the
spe­cies nor subspecies is attributed to an author
in Bruguière (1792), he is deemed the author of
the name (ICZN 1999: Art. 50.1).
Clench (1942) designated the illustration of
the specimen from the Spengler collection illu­
strated in Chemnitz (1788: Pl. 141, fig. 1306) as
representation of the lectotype of C. cedonulli
dominicanus (Pl. 1, Fig. 14). Clench was una­ware
that the figured specimen existed in the ZMUC,
as was Kohn (1968, 1992), but Coo­mans, Mool­
enbeek & Wils (1985) identified and illustrated
it as the lectotype (Figs. 15, 16).
Vink & von Cosel (1985) considered C. c.
dominicanus a valid subspecies of C. cedonulli
Linnaeus in Grenada and the Grena­dines from
Bequia to Carriacou. However, they were unable to examine the lectotype. It conforms more
closely to their description of C. c. cedonulli, the
orange pigment covering most of the shell rather
than being restricted to spiral bands (Kohn, in
prep.). C. c. dominicanus Hwass in Bruguière,
1792 is tentatively concluded to be a junior
synonym of C. c. cedonulli Linnaeus, 1767. If
the more southern Grenadines populations of C.
cedonulli merit subspecies status a different tri­
nomial would be needed.
Conus cinereus Hwass in Bruguière, 1792
Pl. 1, Figs. 17–19
Material examined:
Paralectotype of Conus cinereus: ZMUC-GAS-353.
Remarks
Cernohorsky (1974) recognized the specimen
shown in Figs. 18–19 as a syntype of Hwass’s
Variety C of C. cinereus. It is the specimen shown
in Chemnitz (1788: pl. 142, fig. 1319) (Fig. 17),
one of three figures cited (Hwass in Bruguière,
1792: 673) to illustrate the concept of Variety C,
for which no specimen exists in the Hwass collection (Mermod 1947, Kohn 1992).
Conus cinereus Hwass in Bruguière is a valid
Indo-West Pacific species (Röckel, Korn & Kohn
1995). Its lectotype is a specimen of Hwass’s Variety A from the Hwass collection in the MHNG
designated by Kohn (1968). Because of the prior
lectotype designation, the ZMUC specimen of
Variety C (Figs. 18, 19), originally from the
Moltke collection, is no longer a syntype but is
a paralectotype, according to Art. 73.2.2 of the
ICZN (1999).
Cucullus equestris Röding, 1798
Pl. 2, Figs. 20–22
Material examined:
Lectotype of Cucullus equestris: ZMUC-GAS-354 (= Conus
bandanus Hwass in Bruguière, 1792). [Note: This specimen
is also the lectotype of Cucullus torquatus Röding, 1798.]
Remarks
Peter Friedrich Röding (1767–1846), a little
known 18th Century German naturalist (Dance
1986) prepared a sale catalogue, the Museum
Bolten­ianum (Röding 1798), for the extensive
collection of Dr. Joachim Friedrich Bolten
(1718–1796), a well-known Hamburg physician and member of the Academy of Sciences of
Berlin. Röding (1798) assigned the 157 nominal
species of Conus he listed to the new genus
Cucullus, and he introduced 99 new species
names. Kohn (1992) described the organization of the Museum Bol­tenianum, its treatment
of Cucullus, and its nomenclatural history, and
Dance (1986) dis­cussed the history of the Bolten
collection. Parts of it are now in the Museum
der Natur, Gotha, but Spengler evidently possessed one of the two specimens Röding (1798:
38, No. 474/6) described as C. equestris, and it
is now in the ZMUC. Röding (1798: 38), cited
the illu­stration in Chem­nitz (1788: pl. 138, fig.
1279) of a specimen Chemnitz (1788: 21) stated
to be in the Spengler collection (Pl. 2, Fig. 20),
and Kohn (1975) designated it as the lectotype
(Pl. 2, Figs. 21, 22). It is also the lectotype of C.
torquatus (Röding, 1798), q.v. below. Chemnitz
(1788: 21–22) cited the locality of the lectotype
as “Ost­indischen Meere.”
In addition to applying two species names to
the same specimen, Röding (1798) used the name
C. equestris for two different species-group taxa;
they are thus primary homonyms. The second
(Röding 1798: 46, no. 578/[79]) is a form of C.
type specimens of conus
ammiralis Linnaeus, 1758, of infrasubspecific
rank (Kohn 1992).
Coomans, Moolenbeek & Wils (1985) con­
cluded that the lectotype of C. equestris (Röding)
is an oddly marked specimen of C. bandanus
Hwass in Bruguière, 1792, and Kohn (1992) concurred. C. equestris (Röding, 1798: 38) is thus
concluded to be a junior synonym of C. ban­danus
Hwass in Bruguière, 1792, a widely dis­tributed
species throughout most of the tro­pical IndoPacific region (Röckel, Korn & Kohn 1995).
Cucullus millepunctatus Röding, 1798
Pl. 2, Figs. 23–25
Material examined:
Lectotype of Cucullus millepunctatus: ZMUC-GAS-356 (=
Conus puncticulatus Hwass in Bruguière, 1792).
Remarks
Röding (1798: 47, no. 605/99) cited Gmelin
(1791: sp. 36, “C. leucostictus ε”) and Chemnitz
(1788: pl. 140, fig, 1305) as indications to C.
mille­punctatus. As the latter (Pl. 2, Fig. 23) was
the only published figure cited, Kohn (1975)
de­signated the figured specimen, in the ZMUC,
as lectotype of Conus millepunctatus (Röding,
1798) (Pl. 2, Figs. 24, 25). The specimen pre­
sumably came from the Bolten collection via
Spengler to the ZMUC, although Chemnitz
(1788: 46) cited it as belonging to Spengler at
that time.
As the lectotype of C. millepunctatus (Röding,
1798) is a specimen of the western Atlantic spe­
cies previously described as C. puncticulatus
Hwass in Bruguière, 1792, C. millepunctatus
(Röding, 1798) is a junior syn­onym of C. punc­
ticulatus Hwass in Bruguière, 1792 (Kohn
1992).
Cucullus quadratus Röding, 1798
Pl. 2, Figs. 26–28
Material examined:
Lectotype of Cucullus quatratus: ZMUC-GAS-357 (= Conus
inscriptus Reeve, 1843). [Note: This specimen is also a syntype of C. inscriptus, Reeve, 1843.]
Remarks
After Kohn (1975) designated the cited figure in
Chemnitz (1788: pl. 140, fig. 1300) (reproduced
103
here as Pl. 2, Fig. 26) as a representation of the
lectotype of C. quadratus, Dr Dieter Röckel
iden­tified the specimen in the ZMUC, and Kohn
(1992) designated it as the lectotype (Pl. 2, Figs.
27, 28). The specimen was in Spengler’s collec­
tion after the Bolten sale, although Chemnitz
(1788: 42) listed it as in his possession at that
time. A more recent label indicates that it had
originally come from Peter Forsskål’s expedition
to Arabia in 1761–1767.
In earlier works (e.g. Kohn 1992) I syno­
nymized C. quadratus under C. spurius Gmelin,
1791, a Western Atlantic species. However,
fol­lowing subsequent studies of Indian Ocean
Conus (Kohn 2001, Hylleberg & Kilburn 2002:
p. 47, fig. 47) it has become clear that the lecto­
type of C. quadratus (Röding, 1798) is a spec­
imen of C. inscriptus Reeve, 1843. That species is
generally restricted to continental regions of the
Indian Ocean, from Natal to the southern Red Sea
and eastward to western Thailand (Röckel, Korn
& Kohn 1995), so it could well have come from
Forsskål’s journey.
Although Röding’s name is earlier than
Reeve’s by 46 years, it has not been used as a
valid name in the literature and thus must be
considered a nomen oblitum. C. inscriptus Reeve
has been used as the valid name of the species in
at least 25 works by more than 18 authors during the period 1977–2007; the list is available
at The Conus Biodiversity Website (http://biology. burke.washington.edu/conus). C. inscriptus
Reeve, 1843 is thus the valid name of the species
and a nomen protectum (ICZN, Art. 23.9).
Cucullus summus Röding, 1798
Pl. 2, Figs. 29–31
Material examined:
Lectotype of Cucullus summus: ZMUC-GAS-359 (= Conus
ammiralis Linnaeus, 1758).
Remarks
Kohn (1975) designated fig. 1307 on plate 141 in
Chemnitz (1788) (Pl. 2, Fig. 29) as repre­sentation
of the lectotype of C. summus. Röding (1798)
cited only that Chemnitz figure as an indication
for the nominal species, although the lot of which
it was a part in the Bolten collection comprised
four shells. Subsequently, Filmer (2001) iden­
104
a. j. kohn
tified the figured specimen in the ZMUC; it is
thus the lectotype of C. summus (Röding, 1798)
(Pl. 2, Figs. 30, 31). The spec­imen, originally in
the Bolten collection, was later in Chemnitz’s
collection, then that of King Christian VIII,
which came to the ZMUC in 1848.
The lectotype and only known extant spec­
imen of Conus summus Röding (1798) is clearly
of Conus ammiralis Linnaeus, 1758, and the
name is thus a junior synonym of the latter spe­
cies.
Cucullus torquatus Röding, 1798
Pl. 2, Figs. 20–22
Material examined:
Lectotype of Cucullus torquatus: ZMUC-GAS-354 (= Conus
bandanus Hwass in Bruguière, 1792). [Note: This specimen
is also the lectotype of Cucullus equestris Röding, 1798.]
Remarks
As noted above and in Kohn (1992), the lecto­type
of C. torquatus (Röding, 1798: 38) is the same
specimen as that of C. equestris (Röding, 1798:
38) (pl. 2, Figs. 20–22), making these speciesgroup names objective synonyms. Both are thus
junior synonyms of C. bandanus Hwass in Bru­
guière, 1792 (Kohn 1992).
Conus tenellus Holten, 1802
Pl. 2, Figs. 32–35
Material examined:
Holotype of Conus tenellus Holten, 1802: ZMUC-GAS-360
(= C. nimbosus Hwass in Bruguière, 1792). [Note: This specimen is also the lectotype of C. tenellus Dillwyn, 1817.]
Remarks
Holten (1802) based Conus tenellus solely on the
two figures in Vol. 11 of the Neues Syste­ma­tisches
Conchylien-Cabinet (Chemnitz 1795, pl. 183,
figs. 1782, 1783) (Pl. 2, Figs. 32, 33). Be­cause
this specimen, from the Spengler collec­tion, was
the only one Chemnitz mentioned, and because
Holten (1802: 39) refers to it as “sp. unic.,” it is
the holotype of C. tenellus Holten, 1802 (Kohn
1992) (Pl. 2, Figs. 34, 35).
Because this species had been described a decade earlier by C.H. Hwass (in Bruguière 1792:
732) as C. nimbosus, C. tenellus Holten, 1802 is
a junior synonym of C. nimbosus Hwass in Bru­
guière, 1792 (Kohn 1992).
Conus violaceus Link, 1807
Pl. 2, Figs. 36–38)
Material examined:
Holotype of Conus violaceus: ZMUC-GAS-362 (= C. glans
Hwass in Bruguière, 1792).
Remarks
Heinrich Friedrich Link (1767–1851) was a pro­
fessor of natural history, chemistry and botany at
the universities of Rostock, Breslau, and Berlin
and was the director of the University of Berlin
Botanic Garden. While at Rostock (1792–1811),
Link published the Beschriebung der Natura­
lien-Sammlung der Universität zu Rostock in
1806–1808. The second and third of four parts
(Link 1807) describes the University’s collection of molluscs and other shelled invertebrates.
Link’s enumeration included 75 Recent species
of Conus, for which he introduced 19 new names
(Kohn 1992).
Link applied several of his new names, includ­
ing C. violaceus, to taxa that Gmelin (1791) had
considered varieties of Linnaean species but that
Plate 2: 20. Conus equestris (Röding) and C. torquatus
(Röding) (= C. bandanus Hwass in Bruguière). Illustration
of lectotype reproduced from Chemnitz (1788: pl. 138, fig.
1279). – 21, 22. Conus equestris (Röding) and C. torquatus
(Röding) (= C. bandanus Hwass in Bruguière). Lectotype
(ZMUC-GAS-354). 48x25 mm. – 23. Conus millepunctatus
(Röding) (= C. puncticulatus Hwass in Bruguière). Illustration of lectotype reproduced from Chemnitz (1788: pl. 140,
fig. 1305). – 24, 25. Conus millepunctatus (Röding) (= C.
puncticulatus Hwass in Bruguière). Lectotype (ZMUCGAS-356). 24x15.5 mm. – 26. Conus quadratus (Röding)
(= C. inscriptus Reeve). Illustration of lectotype reproduced
from Chemnitz (1788: pl. 140, fig. 1300). – 27, 28. Conus
quadratus (Röding) (= C. inscriptus Reeve). Lectotype of C.
quadratus and syntype of C. inscriptus (ZMUC-GAS-357).
34x17. – 29. Conus summus (Röding) (= C. ammiralis Linnaeus). Illustration of lectotype reproduced from Chemnitz
(1788: pl. 141, fig. 1307). – 30, 31. Conus summus (Röding)
(= C. ammiralis Linnaeus). Lectotype (ZMUC-GAS-359).
44x24 mm. – 32, 33. Conus tenellus Holten and C. tenellus
Dillwyn (= C. nimbosus Hwass in Bruguière). Illustration of
holotype of C. tenellus Holten and lectotype of C. tenellus
Dillwyn, reproduced from Chemnitz (1795: pl. 183, figs.
1782, 1783). – 34, 35. Conus tenellus Holten (= C. nimbosus
Hwass in Bruguière). Holotype (ZMUC-GAS-360). Also
lectotype of C. tenellus Dillwyn. 34x16 mm. – 36. Conus
violaceus Link (= C. glans Hwass in Bruguière). Illustration
of holotype reproduced from Chemnitz (1788: pl. 143, fig.
1331). – 37, 38. Conus violaceus Link (= C. glans Hwass in
Bruguière). Holotype (ZMUC-GAS-362). 45x20 mm.
type specimens of conus
105
106
a. j. kohn
Link considered distinct species. His entire entry
(Link 1807: 106) is: “C. violaceus. Vielfarbige
K[egelschnecke]. C. Terebellum γ. Linn. Gm. p.
3390. Chemn. Conch. 10. t. 140 [error for 143].
f. 1331. n. 1. Mit C. Terebellum hat sie gar keine
Aehnlichkeit.”
The ZMUC specimen (Pl. 2, Figs. 37, 38)
clearly matches the illustration in Chemnitz
(1788: pl. 143, fig. 13311) (Pl. 2, Fig. 36) cited in
Link (1807), and is the holotype of C. violaceus
(Kohn 1992). The inscription on the shell (Fig.
38), indicates that the holotype derives from the
Moltke collection.
Conus violaceus Link, 1807 is a junior syno­
nym of C. glans Hwass in Bruguière, 1792 (Kohn
1992), and a widely distributed Indo-Pacific spe­
cies (Röckel, Korn & Kohn 1995).
Conus festivus Dillwyn, 1817
Pl. 3, Figs. 39–42
Material examined:
Lectotype of Conus festivus: ZMUC-GAS-363 (= C. pertusus
Hwass in Bruguière, 1792).
Remarks
Lewis Weston Dillwyn (1778–1855) introduced
16 new Conus species names in his Descriptive
Catalogue of Recent Shells, published in 1817,
intended as an update and clarification of Gme­
lin’s 13th edition of the Systema Naturae (Kohn
1992). However, 13 of these names are inad­
vertent introductions. They are of species names
that originated in works now on the Official List
of Rejected Works in Zoology (Hemming 1958),
but that were made available prior to 1817 by
other authors that Dillwyn did not cite. Included
are this and the next species listed, attributed by
Dillwyn to Chemnitz, but under the current rules
of nomenclature they were made available in
Dillwyn (1817).
Dillwyn (1817: 413) cited several descrip­tions
and previously published figures in his description of “C. festivus Chemnitz” (Kohn 1992). The
first of these listed (Chemnitz 1795: 57, pl. 182,
figs. 1770, 1771) (Pl. 3, Figs. 39, 40) shows the
lectotype, from the Spengler collection in the
ZMUC (Pl. 3, Figs. 41, 42) (Kohn 1992).
Prior to Dillwyn, Hwass (in Bruguière, 1792:
673) described this species as C. pertusus (Kohn
1992). C. festivus Dillwyn, 1817 is thus a junior
synonym of C. pertusus Hwass in Bruguière,
1792. C. pertusus is widely distributed through­
out the tropical Indo-Pacific region (Röckel,
Korn & Kohn 1995).
Conus nisus Dillwyn, 1817
Pl. 3, Figs. 43–46
Material examined:
Holotype of Conus nisus: ZMUC-GAS-364 (= C. cinereus
Hwass in Bruguière, 1792).
Remarks
Dillwyn (1817: 388) based his “C. nisus Chem­
nitz” solely on the specimen described and
illu­strated in Chemnitz (1795: 64, pl. 183, figs.
1784, 1785) (Pl. 3, Figs. 43, 44). The illustrated
spec­imen, from the Spengler collection, is in the
ZMUC (Pl. 3, Figs. 45, 46) and is the holotype
of C. nisus Dillwyn (Kohn 1992).
As in the case of C. festivus Dillwyn, Hwass
(in Bruguière 1792: 673) had already described
his C. nisus as C. cinereus (Kohn, 1992). C. nisus
Dillwyn, 1817 is thus a junior synonym of C. cinereus Hwass in Bruguière, 1792. C. cinereus occurs in the western tropical Pacific from south­ern
Japan to New Caledonia and west to Indo­nesia
(Röckel, Korn & Kohn 1995).
Conus tenellus Dillwyn, 1817
Pl. 2, Figs. 32–35
Material examined:
Lectotype of Conus tenellus Dillwyn, 1817: ZMUC-GAS-360
(= C. nimbosus Hwass in Bruguière, 1792). [Note: This specimen is also the holotype of C. tenellus Holten, 1802.]
Remarks
Dillwyn (1817) described C. tenellus, without
reference to Holten (1802). He likewise based
his species on the same cited figures in Chemnitz
(1795) but also referred to a specimen described
by Lamarck. The holotype of C. tenellus Holten,
1802 (Pl. 2, Figs. 34, 35) is thus also the lectotype
of C. tenellus Dillwyn, 1817, and the latter is an
objective primary homonym as well as a junior
synonym of C. tenellus Holten, 1802 (Kohn
1992).
type specimens of conus
107
Conus erythraeensis Reeve, 1843
Pl. 3, Figs. 47–49
a valid species occurring along the continental
margins of the Indian Ocean.
Material examined:
Lectotype of Conus erythraeensis: ZMUC-GAS-365.
Conus tabidus Reeve, 1844
Remarks
In his Conchologia Iconica, the English malaco­
logical author and publisher Lovell Augustus
Reeve (1814–1865) (Petit 2007) described Co­
nus erythraeensis based on specimens from the
collection of F. J. Stainforth (see below) and King
Christian VIII. Reeve attributed the species name
to a manuscript by the Danish zoologist Henrick
Henricksen Beck (1799–1863) whom the King,
while he was still Prince Christian Frederik, had
appointed to manage his collection (Vedelsby
2000).
In their review of the taxonomy of the species,
Coomans, Moolenbeek & Wils (1986) de­sig­
nated a specimen from the King’s collection in
the ZMUC (Pl. 3, Figs. 48, 49) as the lectotype
of C. erythraeensis Reeve. Although the shell is
smaller than the illustration in Reeve (1843: pl.
24, sp. 137; 24.3x14.6 vs. 28x15 mm) (Pl. 3, Fig.
47), its color pattern is similar.
Conus erythraeensis Reeve, 1843 is a valid
species, occurring in the central and southern
Red Sea and the Gulf of Aden (Röckel, Korn &
Kohn 1995).
Conus inscriptus Reeve, 1843
Pl. 2, Figs. 26–28
Material examined:
Paralectotype of Conus inscriptus: ZMUC-GAS-357. [Note:
This specimen is also the lectotype of Cucullus quadratus,
Röding, 1798.]
Remarks
As mentioned above, Reeve (1843) described
Conus inscriptus. Three specimens from the type
lot are in The Na­tural History Museum, London
(nos. 1981052/1–3). Röckel et al. (1995: pl. 38,
fig. 1) designated the first of these as lectotype;
the other two thus became paralectotypes. In his
original description, Reeve (1843: pl. 29, sp. 164)
cited pl. 140, fig. 1300 in Chemnitz (1788) as a
published illustration of his new species. Thus,
the ZMUC specimen (Pl. 2, Figs. 26, 27) is also
a paralectotype of C. inscriptus Reeve, 1843,
Pl. 3, Figs. 50–52
Material examined:
Holotype of Conus tabidus: ZMUC-GAS-366.
Remarks
Reeve (1844) described Conus tabidus based
solely on a single specimen from the collection
of the Reverend Francis John Stainforth (1797–
1866), a rector in London who provided Reeve
with many of the specimens illustrated in that
work (Dance 1986). Stainforth was also an early
philatelist who founded the first stamp collec­tors’
club, which after his death became the Phi­la­telic
Society of London.
The whereabouts of the described specimen
has been generally unknown, but in an undated
note with a specimen in the ZMUC (Pl. 3, Figs.
51, 52), Robert Moolenbeek identified it as pos­
sibly the holotype. Filmer (2001) also noted this
specimen as “possible holotype in ZMUC.” I
examined the unmarked specimen and conclude
that it is almost certainly the figured specimen in
Reeve (1844) (Pl. 3, Fig. 50), based on its size,
shape, and the unusual broad collabral tan streaks
on the last whorl. The latter are lighter than
Reeve’s figure indicates but they are definitely
visible. I thus conclude that the specimen in the
ZMUC (Pl. 3, Figs.51, 52) is the holotype of C.
tabidus. The specimen was likely purchased from
Reeve for the King’s collection, but its original
locality is unknown.
C. tabidus Reeve, 1844 is generally consid­
ered a valid species, occurring along the West
African coast and in the Cape Verde Islands
(Rolán 1991, Cunha et al. 2005).
Conus agrestis Mörch, 1850
Pl. 3, Figs. 53, 54
Material examined:
Paratype of Conus agrestis: ZMUC-GAS-367 (= C. loroisii
Kiener, 1845).
Remarks
In 1850, Otto Andreas Lowson Mörch (1828–
1878), a conchological assistant at the ZMUC
108
a. j. kohn
who wrote extensively on Vermetidae and sup­
plemented his meagre salary by identifying
spec­imens and cataloguing collections for sale,
pub­lished a sale catalogue for the collection of C.
P. Kierulf, Catalogus Conchyliorum quae reli­quit
C. P. Kierulf, who had collected in the Philip­pines
(Dance 1986).
Mörch introduced one new species-group
name in Conus in this work, C. agrestis. The
type specimen was in Kierulf’s collection, but
it was sold at auction in 1850, and its present
where­abouts are unknown. Coomans, Moolenbeek & Wils (1979) translated the description
of C. agres­tis into English and noted that Mörch
also cited a specimen figured by Chemnitz in
the Neues Systematisches Conchylien-Cabinet,
but he cited it incorrectly. However, Coomans,
Moo­lenbeek & Wils (1979) believed that some of
the eight specimens Mörch studied were in the
ZMUC. Jørgen Knudsen and Robert Moolen­
beek then located a specimen from Tranquebar in
the ZMUC they considered to be one studied by
Mörch, and considered it a paratype of C. agrestis
(Pl. 3, Figs. 53, 54). Coomans, Moolen­beek &
Wils (1979) concluded that this specimen represents a form of Conus figulinus loroisii Kiener,
1845. I agree, but with Filmer (2001) consider
C. loroisii a distinct species from C. figulinus
Linnaeus, 1758 (see also Röckel, Korn & Kohn
1995). C. agrestis Mörch, 1850 is thus a junior
synonym of C. loroisii Kiener, 1845.
Conus knudseni Sander, 1982
Pl. 3, Figs. 55, 56
Material examined:
Holotype of Conus knudseni: ZMUC-GAS-368; para­type:
ZMUC-GAS-369 (both = C. sanderi Wils & Moolenbeek,
1979).
Remarks
This and the following species were described
by Dr Finn Sander (1982), then director of the
Bellairs Research Institute, Barbados, and their
disposition is considered together below. A part
of McGill University, Bellairs is the only Ca­
nadian tropical marine research institution. Dr.
Sander was later Administrative Officer of the
Biology Department at Dalhousie University,
Halifax, Nova Scotia, retiring in 2004. He named
C. knudseni in honor of Dr Jørgen Knudsen. It
was based on only three specimens, of which the
holotype (Pl. 3, Figs. 55, 56) and one paratype
are in the ZMUC. All were dredged along the
west coast of Barbados off St. James at a depth
of about 175m.
Conus sorenseni Sander, 1982
Pl. 3, Figs. 57, 58
Material examined:
Holotype of Conus sorenseni: ZMUC-GAS-370; para­type:
ZMUC-GAS-371 (both = C. sanderi Wils & Moolenbeek,
1979).
Remarks
C. sorenseni was named after Mr Ole Sørensen, a
conchologist from Rancho Santa Fe, California.
Like C. knudseni, it was based on only three specimens. Sander (1982) deposited the holotype (Pl.
3, Figs. 57, 58), the only known adult spec­imen,
and one paratype (21x10.3 mm) in the ZMUC.
A subsequent study of additional specimens of
both species by Vink & Sander (1983) reviewed
the taxonomy of C. knudseni and C. sorenseni and
concluded that all specimens vary contin­uously
in shell shape and color pattern with each other
and with C. sanderi Wils & Moolenbeek, 1979.
The three nominal species were described from
the same locality. I agree with the con­clusion of
Plate 3: 39, 40. Conus festivus Dillwyn (= C. pertusus Hwass
in Bruguière). Illustration of lectotype reproduced from
Chemnitz (1795: pl. 182, figs. 1770, 1771). – 41, 42. Conus
festivus Dillwyn (= C. pertusus Hwass in Bruguière). Lectotype (ZMUC-GAS-363). 23x13 mm. – 43, 44. Conus nisus
Dillwyn (= C. cinereus Hwass in Bruguière). Illustration of
holotype reproduced from Chemnitz (1795: pl. 183, figs.
1784, 1785). – 45, 46. Conus nisus Dillwyn (= C. cinereus
Hwass in Bruguière). Holotype (ZMUC-GAS-364). 29x13
mm. – 47. Conus erythraeensis Reeve. Illustration of lectotype reproduced from original description by Reeve (1843: pl.
24, sp. 137). – 48, 49. Conus erythraeensis Reeve. Lectotype
(ZMUC-GAS-365). 24x15 mm. – 50. Conus tabidus Reeve.
Illustration of holotype reproduced from original description
by Reeve (1844: pl. 44, sp. 243). – 51, 52. Conus tabidus
Reeve. Lectotype (ZMUC-GAS-366). 34x19 mm. – 53,
54. Conus agrestis Mörch (= C. loroisii Kiener). Paratype
(ZMUC-GAS-367). 76x49 mm. – 55, 56. Conus knudseni
Sander (= C. sanderi Moolenbeek and Wils). Holotype
(ZMUC-GAS-368). 23x11 mm. – 57, 58. Conus sorenseni
Sander (= C. sanderi Moolenbeek and Wils). Holotype
(ZMUC-GAS-370). 34x17 mm.
type specimens of conus
109
Sander
Sander
Sander
Sander
knudseni
knudseni
sorenseni
sorenseni
8
10 3
1
1
Holotypes
Lectotypes
Paratypes
Syntypes
Paralectotypes None
None
None
None
Chemnitz 1795, pl. 183, f. 1782-1783
Chemnitz 1788, pl. 143, f. 1331, no. 1
Chemnitz 1795, pl. 182, f. 1770, 1771
Chemnitz 1795, pl. 183, f. 1784, 1785
Chemnitz 1795, pl. 183, f. 1782, 1783
None
Chemnitz 1788, pl. 140, f. 1300
None
None
8
10 1
1
1
8
8
3
0
1
Number of Species Number of Type Specimens
= sanderi Wils & Moolenbeek, 1979
= sanderi Wils & Moolenbeek, 1979
= sanderi Wils & Moolenbeek, 1979
= sanderi Wils & Moolenbeek, 1979
= nimbosus Hwass in Bruguière, 1792
= glans Hwass in Bruguière, 1792
= pertusus Hwass in Bruguière, 1792
= cinereus Hwass in Bruguière, 1792
= nimbosus Hwass in Bruguière, 1792
valid
valid
valid = loroisi Kiener, 1845
= cedonulli cedonulli Linnaeus, 1767
Chemnitz 1788, pl. 141, f. 1306 valid
Chemnitz 1788, pl. 142, f. 1319 = bandanus Hwass in Bruguière, 1792
Chemnitz 1788, pl. 138, f. 1279 = puncticulatus Hwass in Bruguière, 1792Chemnitz 1788, pl. 140, f. 1305
= inscriptus Reeve, 1843
Chemnitz 1788, pl. 140, f. 1300
= ammiralis Linnaeus, 1758
Chemnitz 1788, pl. 141, f. 1307
= bandanus Hwass in Bruguière, 1792
Chemnitz 1788, pl. 138, f. 1279
None
Chemnitz 1788, pl. 141, f. 1310
Martini 1773, pl. 59, f. 655 Figure Prior to Species Publication
Identification of collections: B, Bolten; C, Chemnitz; CVIII, King Christian VIII; M, Moltke; S, Spengler.
Totals
23 211 202,3,4
1
Two of the paratypes are of species whose holotypes are also present.
2
The same specimen is the lectotype of C. equestris (Röding, 1798) and C. torquatus (Röding, 1798).
3
The same specimen is the lectotype of C. quadratus (Röding, 1798), an unused senior synonym, and a syntype of C. inscriptus Reeve, 1843.
4
The same specimen is the holotype of C. tenellus Holten, 1802 and the lectotype of C. tenellus Dillwyn, 1817.
Entries in Table
GAS-368
GAS-369
GAS-370
GAS-371
GAS-360
GAS-362
GAS-363
GAS-364
GAS-360
GAS-365
GAS-357
GAS-366
GAS-367
GAS-352
GAS-353
GAS-354
GAS-356
GAS-357
GAS-359
GAS-354
Table 1, Cont.
1982
1982
1982
1982
Holotype Paratype Holotype Paratype Holotype4 Holotype Lectotype Holotype Lectotype4
Lectotype Paralectotype3
Holotype Paratype 1802
1807
1817
1817
1817
1843
1843
1844
1850
Holten
Link
Dillwyn
Dillwyn
Dillwyn
Reeve
Reeve
Reeve
Mörch
tenellus
violaceus festivus
nisus
tenellus
erythraeensis
inscriptus
tabidus
agrestis
ZMUC No.Current Species Name
Holotype
GAS-114 valid
Holotype
GAS-351 = mappa (Lightfoot, 1786)
Lectotype GAS-168 = centurio Born, 1778
Date Type
Lectotype Paralectotype
Lectotype2
Lectotype
Lectotype3
Lectotype Lectotype2 Author
gloriamaris Chemnitz 1777
solidus
Gmelin
1791
tribunus Gmelin
1791
cedonulli
dominicanus Hw. in Brug.1792
cinereus Hw. in Brug.1792
equestris Röding
1798
millepunctatus Röding
1798
quadratus
Röding
1798
summus
Röding
1798
torquatus
Röding
1798
Species Table 1. Type specimens of Conus in the Zoological Museum of the University of Copenhagen.
S
M
S
S
S
CVIII
C→B→S
CVIII(?)
S
M
S → B (?)
S(?) → B → S
C→B→S
B → C(?) → CVIII
S → B (?)
M
M → CVIII
S
Prior History of Specimen
110
a. j. kohn
type specimens of conus
Vink & Sander (1983) that all are conspecific
and consider C. sorenseni Sander, 1982 and C.
knudseni Sander, 1982 to be junior synonyms of
C. sanderi Wils & Moolenbeek, 1979.
DISCUSSION AND CONCLUSIONS
The ZMUC holds 20 type specimens of Conus.
Of these, 16 are primary types – holotypes and
lectotypes – that served as the basis for 19 species of this hyperdiverse marine gastropod genus,
and several have resided in the Museum since
the Museum Moltkianum came to the University
nearly 200 years ago. Their pub­lication dates
extend back even farther, to Chemnitz’s descrip­
tion of the famous Conus gloriamaris in 1777,
and they are summarized chronologically in
Table 1.
The original ownership of some specimens
is uncertain, but four of the 18th and early 19th
Century specimens derive from the collections
of Adam and Joachim Moltke that formed a
large part of the original Museum, and nine are
from the Lorenz Spengler collection; some of the
latter had belonged to J. F. Bolten. The rest had
be­longed to J. H. Chemnitz and King Christian
VIII (Table 1).
These shells are identifiable because they
were described and well illustrated in the Syste­
matisches Conchylien-Cabinet of Martini and
Chemnitz (1773–1795). Eight of the primary
type specimens served as the basis of 10 species
described as new in the 18th Century. Only one
of these, C. gloriamaris Chemnitz, is presently
con­sidered a valid species. Six of the nine 19thCentury species whose type specimens are in the
ZMUC were also illustrated by Chemnitz (1788–
1795). One of the 18th-Century lectotypes is of an
unused senior synonym but is also a syntype of
C. inscriptus Reeve, described in 1843. It and the
little known C. tabidus Reeve, 1844 are the only
19th-Century species presently considered valid.
The only 20th-Century Conus types in the ZMUC
are C. knudseni and C. sorenseni, described by
Sander (1982) but shortly thereafter sunk in syn­
onymy (Vink & Sander 1983).
Three new conclusions concerning type spec­
imens and taxonomy result from this study: The
previously designated lectotype of C. amadis
111
Gmelin, 1791 (Kohn 1966, 1992) loses its lecto­
type status and is replaced by a representation
of the lectotype; the lectotype of C. cedonulli
domi­nicanus Hwass in Bruguière, 1792 is not
distin­guishable from C. cedonulli cedonulli Linnaeus, 1767; and the existence of the holotype
of C. tabidus Reeve, 1844 was not previously
rec­ognized.
Although the original names of only five of
the 16 ZMUC type specimens of Conus remain
valid, the type collection is particularly valuable.
Study of the specimens it comprises enabled the
taxonomic results reported here, and these in turn
led to solving problems of the correct names of
species in this highly diverse and ecologically
important group of marine animals.
DEDICATION
Significant anniversaries of several notable events
in Scandinavian natural history and sy­stematics
occurred in 2007–2008. These in­clude the tercentenary of Linnaeus’s birth and the 250th anniversary of his Systema Naturae (tenth edi­tion)
and, on a more somber note, the 280th anni­versary
of the Copenhagen fire that destroyed early 18th
Century museum collections of ani­mals. This
paper is dedicated to Jørgen Knudsen on a happier occasion, his 90th birthday on 6 March 2008.
Another anniversary, likely signi­ficant only to the
author, is that just 50 years ago I first visited the
ZMUC to study the type spec­imens of the great
gastropod genus Conus, many two centuries old
and carefully held in the Muse­um, then in Krystalgade. It was then I first met Jørgen Knudsen.
He was most helpful in finding those ancient
specimens that still anchor many familiar species
in Conus, the largest genus of marine animals
and one of the taxonomically most vexing. I still
value Jørgen’s generous help half a century ago,
and his insights into mollu­scan functional morphology have been most pro­fitable to me over
the years since.
ACKNOWLEDGEMENTS
NSF Grant 0316338, including a CETAF supple­
ment, supported this study. In addition to Jørgen
112
a. j. kohn
Knudsen’s help in 1958, I thank Antonia Ve­
delsby, Kathe Jensen and Ole Tendal for their
help facilitating my visits to the ZMUC in 2007
and 2008, David Reid for pointing out the du­
bious status of the lectotype of C. amadis and
for other valuable comments on the manuscript,
An­to­nia Vedelsby and Danny Eibye-Jacobsen for
many helpful suggestions on the manuscript, and
Trevor Anderson and Joshua Kubo for assistance
with the illustrations.
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