<oological Journal ofthe Linnean Society (1991), 103: 145-195. With 29 figures
The Linnaean fish collection in the Zoological
Museum of the University of Uppsala
ALWYNE WHEELER
Epping Forest Conservation Centre, High Beach, Loughton, Essex IGlO 4AF
Received December 1990, accepteddfor publication May 1991
Specimens of fishes preserved in the Zoological Museum, University of Uppsala, which are believed
to have been examined by Linnaeus, are listed. Most of these were originally given to the University
in several donations by benefactors of the Academy and were described by Linnaeus in dissertations
defended by students. Some specimens, however, are believed to have originated from Linnaeus’s
own collection. Many of the specimens have type status and this is discussed together with notes on
other surviving Linnaean fish specimens.
KEY WORDS:-Linnaeus
-
fishes - type specimens - Uppsala.
CONTENTS
Introduction . . . . . . . . . . . . . .
Earlier accounts of the collection . . . . . . . . .
The donations to the collection. . . . . . . . . .
Present disposition of the collection
. . . . . . . .
Type specimens of Linnaean species . . . . . . . .
Catalogue of Linnaean specimens in the Zoological Institute Uppsala
Acknowledgements
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References
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INTRODUCTION
This is the third paper which lists and discusses specimens of fish in collections
which were studied by Carl Linnaeus (1707-1778). The first of the series
(Fernholm & Wheeler, 1983) described specimens in the Swedish Museum of
Natural History; the second (Wheeler, 1985) gave an account of the history of
the collection and listed the fishes now preserved in the Linnean Society of
London. The present paper is concerned with the fish specimens in the
Zoological Museum of the University of Uppsala, and includes much of the
earliest material worked on by Linnaeus.
Other studies related to these enumerations of Linnaeus’s collections include
that of the Gronovius collection, the majority of which is now preserved in the
British Museum (Natural History) (Wheeler, 1958). Other, alcohol-preserved
fishes from the Gronovius collection exist in the Zoological Museum,
Copenhagen, and were described in a later paper (Wheeler, 1989). In addition,
I45
0024-4082/91/100145 f 5 1 003.00/0
0 1991 The Linnean Society of London
146
A. WHEELER
Figure 1. Labels in C. P. Thunberg’s autograph are usually the earliest labels on the Uppsala
specimens. They date from the 1780s and relate to the various donations to the collection.
the literature cited by Linnaeus in the tenth and twelfth editions of the Systema
Naturae (Linnaeus, 1758, 1766) was identified and analysed by Wheeler (1979).
The Uppsala collection is ostensibly better documented than either the
Stockholm or London collections of Linnaean fishes because several authors have
published lists of the material contained in it. As early as 1787, Carl Peter
Thunberg, Linnaeus’s eventual successor at the University, published extensive
lists of the material received in several donations to the museum, and made
praiseworthy attempts to curate the material by labelling it fully (Fig. 1). Einar
Lonnberg (1896) published a detailed catalogue of the collection of vertebrate
animals in the Zoological Museum and identified type specimens where possible.
More recently, Ake Holm (1957) prepared and published a complete list of the
zoological material from Linnaeus’s time preserved in the Zoological Museum of
Uppsala University, with a detailed account of the history of the collection.
Each of these analyses of the collection was presented under the sequence by
which the donation was received by the museum, thus maintaining curatorial
integrity, but resulted in the documents having a greater historical than
zoological relevance. The present catalogue of the fishes in the collection is
LINNAEAN FISHES IN UPPSALA
147
arranged systematically, following Nelson ( 1984) for higher categories, and by
family; it is thus slanted towards the ichthyological user rather than the
historian.
EARLIER ACCOUNTS OF THE COLLECTION
As mentioned above, several authors have published lists of the specimens
contained in the Zoological Museum of the University of Uppsala. The first to
do so was Carl Peter Thunberg (1743-1828) who succeeded Linnaeus’s son as
professor of medicine and botany in the University of Uppsala. Thunberg, who
early in life made extensive travels in southern South Africa, Java, Japan and Sri
Lanka, settled in Uppsala to publish a four volume account of his travels
(Thunberg, 1788-93) and several botanical works as well as large numbers of
short papers mostly as academic dissertations describing new taxa and listing
collections of material in the University. This he did without apparently being
aware of developments in zoology and botany that extended these disciplines
beyond the established Linnaean framework. He did, however, recognize that
the Linnaean collections merited curation and appears to have identified,
labelled and catalogued as many of Linnaeus’s specimens as he could locate. He
is also alleged to have substituted better specimens for those in poor condition in
the collection (Wallin, 1985). He has, however, to be credited with a serious
attempt to recognize and document the genuine early Linnaean material in
Uppsala, and Thunberg’s labels on the specimens are a guarantee that the
specimen is at least of eighteenth century origin.
In a published account of the collection in the form of a dissertation defended
by Fredrik Wilhelm Radloff, Thunberg ( 1787) listed the zoological specimens
received by the Academy at Uppsala in a series of donations by benefactors of
the University. The specimens were listed in order of receipt of the donations
between 1744 and 1749 with, in some cases footnotes identifying the specimens
or commenting on their preservation. Parts of Thunberg’s own huge collection
was also listed as donations in 1775 (Thunberg, 1787), and in a series of
Appendixes by Thunberg (1791, 1794, 1798, 1808, 18’18) all of which were
presented as student dissertations. A further listing of material by Thunberg
quoted by Holm (1957) was of the donation by King Gustav IV Adolf to the
University of Uppsala in 1803. Although the donation was clearly post-Linnaean
it included material which had been described by Linnaeus (1764b) in his
account of the collection of Queen Louisa Ulrika. This donation comprised
mostly invertebrates but contained a few vertebrates, including fishes, from the
collection of King Adolf Fredrik in the royal castle of Ulriksdal (see Fernholm &
Wheeler, 1983, for an account of these royal collections).
The arrangement by Thunberg of these catalogues of the collection in
sequence of donations to the University museum was reflected in later
publications on the Linnaean collection, and indeed in the present-day
arrangement of specimens on the shelves in the Uppsala collection. There is,
however, a manuscript catalogue by Thunberg “General-Catalogue p5 Kongel.
Akademiens i Upsala, Natural-samlingar,” begun in 1828 which lists the
holdings of the museum in contemporary systematic order. This also gives the
source of the specimen referring to the donor by abbreviated names (e.g.
Th. = Thunberg, A1 = Alstromer-Linnk-see below), and in cases where there
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A. WHEELER
are several specimens of a taxon identifying each specimen, and thus its source,
with a Greek letter. This is a most valuable indication of the existence of
specimens at this date, and of their origin according to Thunberg.
In the late nineteenth century Einar Lonnberg (1896) published an annotated
catalogue of the vertebrate specimens in the Zoological Museum which had been
associated with, or which were contemporary to Linnaeus. This catalogue was
prefaced by a short historical account of the donations made to the Museum and
some discussion of the type status of the specimens listed. I t was again produced
in the order of the receipt of the donations which, while retaining an historical
perspective, made retrieval of information of taxonomic value extremely
difficult. I t was also a product of its period in that taxonomic practice reflected
the Strickland Code and for this reason some of Lonnberg’s conclusions are
difficult to understand in the present context. It is also true that the presentation
of Lonnberg’s paper is in places ambiguous due to its typographic lay-out, and it
is unfortunate that the author who, with great courtesy published his results in
English, was sometimes less than clear in his meaning. Nevertheless Lonnberg’s
work is the basic critical catalogue for the collection of vertebrates in the
Linnaean Museum at Uppsala.
More recently, the whole of the collection of specimens in the zoological
collection of the university, which date from Linnaeus’s time, have been listed by
Holm (1957), who in addition gave a detailed historical account of the material.
Holm listed the specimens from the various donations (so again the presentation
is historically biased) but relates them to their original names and numbers in
the published dissertations and gives short notes on the existence and method of
preservation of the specimen. He also indicates the putative type status of the
specimens, in the case of the fishes deriving his information mostly from
Lonnberg, but also making use of an unpublished manuscript by 0. Olofsson in
the Zoological Institute at Uppsala.
Holm also reproduced in tabular form the manuscript catalogue of the
Uppsala collection which was compiled within two years of Linnaeus’s death
between 13 September 1780 and 24 March 1781. This listing is critical in
bridging the temporal gap between the Linnaean dissertation in which the
specimens were described and the published catalogues of the collection by
Thunberg (1 787), before Thunberg had extensively curated the collection.
THE DONATIONS TO THE COLLECTION
As already noted, the catalogues of the Uppsala zoological collection
published by Thunberg were arranged in the sequence of donations made to the
university. As far as the fishes are concerned there are four such donations which
are relevant although the two earliest received are the most important and
contain a high proportion of type material.
Donatio Adolphi Friderici
In 1745 the Crown Prince Adolf Fredrik (King from 1751) presented a
collection of animals to Uppsala University. These were described by Linnaeus
(1746a) in a dissertation defended by Laurent Balk under the title Museum
Adolpho-Fridm’cianum.The dissertation was reprinted in the “Linnaeus edition” of
the Amoenitates Academicae (Linnaeus, 1749a) with the running title of “Museum
LINNAEAN FISHES IN UPPSALA
149
Princeps” and this edition, rather than the original printing was the one
generally cited by Linnaeus (1758), usually under the abbreviated title of Amoen.
acad. Elsewhere, as in Linnaeus (1756), he referred to this work by an
abbreviation of the running title M . Pr. or M . princ.
It is important to distingush between references to this publication and to the
later published descriptive account of the collection of King Adolf Fredrik
(Linnaeus, 1754a, 1764a). This was published in two volumes under the title
Museum Adolphi Frederici, the first volume in folio, sumptuously illustrated with
copper-plate engravings, and printed in Latin and Swedish double column,
while the second volume was published in octavo format, in Latin and without
illustration and was issued with the catalogue of the collection of Queen Lovisa
Ulrica, Museum Lodovicae Ulricae (Linnaeus, 1764b). Some species were
represented in the collection of Adolf Fredrik in duplicate or by a series of
specimens. In several cases a specimen was presented to Uppsala in 1745 to be
described by Linnaeus (1746a, 1749a) while the remainder of the series of the
taxon stayed in the royal collection to be described by Linnaeus (1754a, 1764a).
With time the collection of the King became part of the holdings of the
Naturhistoriska riksmuseet, Stockholm (Fernholm & Wheeler, 1983) and many
specimens are still preserved there. In some cases, therefore, specimens which
may be presumed to have been originally part of the same lot in the royal
collection are today found both in Uppsala and in Stockholm (examples are:
Chaetodon saxatilis [ = Abudefduf saxatilis], Chaetodon capistratus, and Cyprinus auratus
[ = Carassius auratus], see Fernholm & Wheeler ( 1983) and this catalogue).
Twenty-one species of fishes were described in Balk’s dissertation (Linnaeus,
1746a, 1749a); 17 of these were recognized by Thunberg (1787) but only 14
were listed as being present in the collection by Lonnberg (1896) and Holm
( 1957).
Donatio Magni Lagerstroem
In 1748 and in 1750 the University of Uppsala received donations from
Magnus Lagerstrom, Councillor of Commerce and a director of the Swedish
East India Company. These were described by Linnaeus (1754b) in a
dissertation Specimen Academicum, sistens Chinensia Lagerstromiana . . ., defended by
Johann Laurens Odhelius, in which 12 species of fishes were described. This
dissertation was referred to by Linnaeus ( 1758) in its original printing under the
abbreviated title Chin. Lagerstr., but in the twelfth edition of the Systema Naturae
(Linnaeus, 1766) additional references are made to the reprinted edition,
Amoenitates Academicae (Linnaeus, 1759).
In 1787 Thunberg listed nine species from this donation, one of which (Gobius
pectinirostris) was represented by two specimens, the second of which may be the
missing Gobius eleotris. Lonnberg (1896) listed only eight taxa from this donation
including the possible specimen of Gobius eleotris, but omitting the specimens of
Cyprinus auratus and Balistes vetula which Thunberg had included. Both these taxa
were included in the list published by Holm (1957) and this author recognizes a
total of ten taxa from the Lagerstrom donation, one of which is a dry mounted
skin.
Several of these fishes are of oriental origin and are presumed to have come to
Lagerstrom through his position as Director of the East India Company, or were
collected in China by him personally. Lagerstrom was instrumental in placing
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A. WHEELER
Pehr Osbeck (1723-1805) as chaplin on the trading vessel Prim Curl of the
Swedish East India Company.
Osbeck’s voyage lasted from November 1750 to June 1752 and during its
course he made observations on the natural history of Spain near Cadiz, Java,
Canton (which was the destination of the trading voyage) and at the Atlantic
island of Ascension. Judging from some of the oceanic fishes he reported in his
account of the voyage he also collected while the ship was at sea (Osbeck, 1757).
Because of the apparent conflict of dates with Lagerstrom donations being
made to the University in 1748 and 1750 and Osbeck’s return in 1752 it seems
on this evidence that none of the Asiatic material in the collection could have
been collected by Osbeck. However, for reasons advanced below I am now
convinced that some of the Uppsala specimens are those collected by Osbeck and
that Thunberg and later authors were in error either in the dating of the
Lagerstrom donation, or that there was another post-June 1752 donation of
Osbeck material which has not previously been recognized.
In this present listing of Uppsala fishes there are six specimens of fishes that
were also listed by Osbeck (1757). These are Clupea thrissa, C. mystus, Lophius
histrio, Gobius pectinirostris, G . eleotris and Balistes vetula. Two other taxa were
named in the English translation of Osbeck’s voyage by J. R. Forster (1771), viz.
Balistes monoceros and Tetraodon ocellatus. The first six were described in Odhelius’s
dissertation Specimen Academicurn s i s t m Chinensia Lagerstromiana (Linnaeus, 1754b)
and attributed by Thunberg to the Lagerstrom donations of 1748 and 1750. The
last two were attributed to the Alstromer-Linnaeus donations of 1749 and later.
It seems to me to be too great a coincidence that even six of the fishes collected
by Osbeck during his voyage should also have been present in the Lagerstom
donations made before he had sailed in 1750. From the Osbeck-Linnaeus
correspondence (and the detailed analysis of it in relation to the plant collection
by Hansen & Maule, 1973) we know that Osbeck had sent some of his collection
including fishes to Lagerstrom before 26 July 1752, and that Lagerstrom had
forwarded the plants (and possibly the animals) to Linnaeus (the latter speaks of
“collections” only) by 10 August. Whether Linnaeus had the fish specimens or
not, they were evidently in the University collection before 1754 when
Odhelius’s dissertation was defended. I t seems to me that there is a very strong
presumption that these six, and probably the additional two specimens, were all
Osbeck material. This means that most of them have a claim to being type
material.
Donation Jonae Alstromer nec non Caroli a Linnt
This heading was used by Thunberg (1787) to include the gift to the
University of natural history specimens by Jonas Alstromer, a Councillor of
Commerce, in 1749. It also includes miscellaneous material added to the
collection by Linnaeus at various times. In many respects this set of specimens is
the least satisfactory of the Uppsala collection because it is difficult to establish
the source, or the history, of most of the specimens. A few are undoubtedly type
specimens of Linnaean species, their standing confirmed by accurate descriptions
by Linnaeus and their rarity in contemporary collections. Many are unlikely to
be type specimens, or at least cannot be proved to be so, and where there are
LINNAEAN FISHES IN UPPSALA
151
specimens of the same taxon in other collections which are more probably types
it seems pointless to strive to find proof of their original status.
Much of this donation comprises Swedish, or at least Palaearctic, species
and is made up of the material which Thunberg could not attribute
unequivocably to any of the recorded donations (there were several other
important donations in addition to those discussed above which did not contain
fish specimens). Some of this material probably originated with Linnaeus-at
least one, the type specimen of Cyprinus cultratus ( = Pelecus cultratus) is known to
have been Linnaeus’s as it was given to him during his journey through Skine
(Linnaeus, 175 1), and Linnaeus may well have placed alcohol-preserved
specimens in the university museum for several reasons including reducing the
fire risk to his library and dry collections. Some of this material may have been
the ‘wet collection’ which was included in the inventory made after the death of
the younger Linnaeus in 1783 by J. G. Acrel. This included an unknown
number of fishes in spirits of wine, although a later list published by C. G. Myrin
(1833) lists “Pisces-in spiritu vini, diversis lagenulis exepti . . .33”. According to
Lowegren (1952) the spirit-preserved material was not sent to London with the
Linnaean library and the natural history collections. This being so, it seems more
probable that it would have been transferrred to the university museum (if it was
not already there) than merely being destroyed.
Many of the specimens in the Alstromer-Linnk donation lack labels other than
those provided by Thunberg in his curation of the collection in the 1780s. It
seems probable that Thunberg, faced with a considerable number of specimens
without labels, was able to relate some to the donations of the Crown Prince
Adolf Fredrik and Magnus Lagerstrom (and some others) by a process of
elimination. The remainder he appears to have labelled and included within this
heterogenous “donation”. As Thunberg provided many of the identifications it is
not proven that these were the names used by Linnaeus and misidentifications
are not necessarily the responsibility of Linnaeus.
Other donations
Two other important collections presented to the university museum
contained a small number of fish specimens. One of these was the Donatio Claudii
Grill of 1746 which contained mostly reptiles but included one fish “Gymnotus
carapo”. Claes Grill was a director of the Swedish East India Company but his
donation comprised animals from Surinam. They were described by Linnaeus in
a dissertation published in 1748 under the title Surinamensia Grilliana (Linnaeus,
1748a) and defended by P. Sundius, which was later reprinted in Amoenitates
Academzcae (Linnaeus, 1 749b). The specimen of Gymnotus was said to be more than
3 feet in length and was listed as missing from the collection by Lonnberg (1896).
The second donation to contain fish specimens some of which were possibly
associated with Linnaeus was the Donatio Gustavi Adolphi of 1803, which was
mentioned earlier (p. 147). Several fishes, or parts of fishes were listed by
Thunberg (as cited by Holm, 1957), viz. Diodon hystrix p, Ostracion cubicus,
Ballistes maculatus, Squalus Acanthias p, four lots of Squali mandibulae, an enigmatic
entry for “Dentes”, and four Pristis rostra. T h e Diodon, Ostracion and Squalus
acanthias were recognized in the Uppsala collection by Holm ( 1957). One of them
may have type status, the others do not, and their connection with Linnaeus is
tenuous.
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Figure 2. Three specimens from the Uppsala Linnaeus collection to show different methods of
preservation and storage jars. Left, Luphiuc vespertilio (ZIU 163) specimen dry, mounted. Centre,
Mywine glutinosa (ZIU 222) specimen in alcohol, jar sealed with glass plate and oxbladder. Right,
Gadus Lo& (ZIU 178), specimen hanging by a thread; jar sealed with glass plate and red sealing wax.
(This jar has not been opened since C. P. Thunberg’s time.) Note: all these specimens have
Thunberg labels.
PRESENT DISPOSITION OF THE COLLECTION
The collection is now the property of the Zoological Institute of the University
of Uppsala. A small number of fish specimens are preserved in the Museum of
the Zoological Institute but the majority of the specimens are displayed in
cabinets in the Linnaean Museum in the Old Botanical Garden in Uppsala, the
official residence occupied by Linnaeus as Professor of Medicine and Botany.
They have been in this building since 1939 and are arranged by order of
donation in glass-fronted cabinets (with a recently-added inner unbreakable
security screen).
Much of the collection is preserved in alcohol and stored in eighteenth century
glassware; a few specimens are preserved as dry or stuffed skins mounted on
wooden plinths by means of wire supports (Fig. 2). The glass cylinders are handblown and thus often uneven in thickness; this together with their age means that
they are very fragile and if handled when empty tend to suddenly craze. They
have a slightly narrowed neck, the top flaring out with a ground surface to the
edge. The tops are sealed with a glass disc in some cases held in place with oxbladder tied tightly with twine, in others the glass is held in place by red sealingwax (Fig. 1). The specimens are often suspended from the lower jaw by twine
attached to the inside of the lid. Labels are mostly small rectangles of paper with
ink inscription, stuck to the foot of the glass container or to the wood mount.
Some labels are attached near the neck of the bottle, Most of the labels are in the
LINNAEAN FISHES IN UPPSALA
153
hand of C. P. Thunberg (see Figs 1, 2). These labels are referred to as ‘original
label’ in the catalogue of the collection; no labels are in Linnaeus’s hand. If
Linnaean labels existed then they may have been removed by Thunberg as part
o f a curatorial tidying-up. Possibly most of them had no Linnaean label, or at
best only a binomen-Linnaeus rarely labelled his zoological specimens with
donor or locality data. Because of this there is an obvious risk of error between
their description by Linnaeus and the curation of the collection by Thunberg,
although a list of the contents of the Museum was prepared in 1780 and the
specimens were subsequently described by Thunberg ( 1 787).
TYPE SPECIMENS OF LINNAEAN SPECIES
Modern authors are conditioned to expect that proposals of new taxa will be
accompanied by a detailed description of the specimen or specimens available for
study and illustration of the animal. With this background reference to the
published source of Linnaean names frequently causes misunderstanding because
the name is usually based upon a short sentence in telegraphic Latin supported
by one or more citations of earlier publications. This is particularly the case with
the Systema Naturue (Linnaeus, 1758, 1766). In both tenth and twelfth editions of
this work the species names appear in the left-hand margin of the page (beneath
a heading of the generic and class or ordinal name), the species are numbered
within the genus serratim and each distinguishing sentence commences with the
initial letter of the genus name. Recent authors usually take this sentence to be
the ‘description’ of the species, and not infrequently are disparaging in their
comments on its brevity and lack of precision. This, however, is a reflection of
their lack of understanding of Linnaeus’s methods.
The Systema Naturae should really be seen as an ‘index’ to the units of the
natural world given names in the binominal method. Against each marginal
trivial or species epithet is a sentence of diagnosis of the features which
distinguish that taxon from others recognized by Linnaeus. This diagnosis is not
a description although Linnaeus regarded it as a sufficient ‘description’ of the
taxon to allow identification to species within the contemporary context. I n most
entries in the work the diagnosis is followed by one or several references to earlier
publications (the identification of these are discussed below). It is only by
reference to these earlier published works that the details of the morphology, or
an illustration can be established. The description that a modern author would
expect has therefore to be culled from the early literature, some of which is
extremely difficult to obtain.
As a consequence, an appreciation of what Linnaeus understood by the name
of a species can only be revealed by examining all the references he cited. Some
of these will lead to descriptions of a specimen or specimens, a few of which will
be as detailed as modern descriptions; others simply refer to a figure in a
publication which Linnaeus regarded as adequate for identification. As a result
of the state of the art of engraving before 1758 (and in some cases for a century
afterwards) the illustrations cited may be no more than crude wood cuts,
although others attained a high standard with copper plate engraving.
It is clear, therefore, that where the Linnaean diagnosis fails to give adequate
details for identification for modern purposes, consideration must be given to the
literary or illustrative sources Linnaeus cited. These number between one and
five, although occasionally there is no citation of literature. I n the latter case
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there are several possible causes. Mostly, these names rely on a specimen
available to Linnaeus which had not been previously formally named, e.g. Gobius
anguillaris (Linnaeus, 1758: 264). I n a few cases the description was based on a
specimen in another collection, e.g. Chaetodon punctatus (Linnaeus, 1758: 273)
“Mus. Ac. Holmens.” ( = the museum of the Academy of Sciences, Stockholm).
In others, no source is cited by Linnaeus in the tenth edition of the System
Naturae, but in the twelfth edition a source is cited, e.g. Gadus mediterraneus
Linnaeus (1758: 255) which is attributed to Mus. Ad. Fr. 2 . p. 60 in Linnaeus
( 1766: 441), this being the second volume of the Museum Adolphi Friderici which
was not published until 1764 (although it was in manuscript,for several years
before that).
Where several literary sources are cited it is necessary to consult these to
establish the concept that Linnaeus had of the species. For fishes citations are
usually given in a sequence commencing with Linnaeus’s own work, then
reference to Artedi, then to Gronovius. This was not an inflexible sequence but
was normally followed. It represents the ‘rankings’ applied by Linnaeus (1 758:
24 1) for the “Ichthyologi Theoretici”, Artedi, himself, Gronovius and
Hasselqvist. This hierarchy represented four authors who consistently gave
detailed descriptions supplemented by good quality illustrations for some taxa by
Linnaeus and Gronovius. In other zoological groups, especially Insecta
Lepidoptera and Vermes Mollusca, illustrated books were of special importance
and form the majority of the literature cited.
In identifying type material of Linnaean animals there must be a structured
sequence of preference. In some cases a name is based on a specimen (sometimes
specimens) examined by Linnaeus for the purposes of a description in a
publication. Clearly these must be regarded as in the front rank for type
designation and, where they can be positively identified, take precedence over
literary references. These can be termed primary types.
A second category arises from specimens which still exist and which were the
subject of description by an author who was cited by Linnaeus in the naming of
the species. Among fishes this is particularly relevant to the Gronovius collection,
a large part of which is preserved in the British Museum (Natural History) and a
small, but important, part in the Zoological Museum, Copenhagen (Wheeler,
1958, 1989).
Among insects there is extant material from De Geer’s collection in Uppsala,
and much of Clerck’s collection of Arachnida also exists (but Clerck employed
binominal nomenclature for his specimens contemporary with Linnaeus but
independently). Where no material exists which can be shown to have been
examined by Linnaeus, but specimens described by another author whose
description was cited are in existence, then these must have validity as type
material. They can be regarded as secondary types.
The third level of material contributing to the typification of Linnaean species
is the literary and artistic references cited in the definition of the species. These
tertiary sources are clearly of minor significance compared with the primary and
secondary sources, when specimens from these exist, but they are nevertheless
important. Experience of this literature shows that it is far more complicated
than might be expected. In the first place, Linnaeus cited books and some serial
publications in a cryptic binary fashion. Many of these abbreviations would be
unintelligible to anyone without a knowledge of eighteenth century literature in
natural history, were it not for the existence of glosses identifying the sources,
LINNAEAN FISHES IN UPPSALA
155
for fishes (Wheeler, 1979), for Mollusca (Boss, 1988), and for all zoology
(Wheeler & Heller, in press). For some early works, simply identifying the title is
not sufficient and it is necessary to check that the edition referred to is that which
Linnaeus cited. This can be verified often by the correspondence of page or plate
numbers but as a last resort reference to the Linnaean library held in the
Linnean Society of London will be required.
The most frequently cited published work in fishes was Artedi (1 738), and this
is unique in every sense. The life and work of Artedi have been discussed by
Lonnberg (1905) and Wheeler (1961, 1987). Artedi’s lchthyologia (1738) which
was edited by Linnaeus, after the author’s death in 1735, comprises five sections,
the Bibliotheca Ichthyologica in which all the significant existing ichthyological
literature is reviewed, the Philosophia lchthyologia in which the framework for the
classification of fishes is discussed, the Genera Piscium which lists all known genera
and species, the Synonymia nominum Piscium where he established detailed
synonymies for the species, and the Descriptiones specierum piscium in which he gave
descriptions of a number of species. The last three parts were extensively cited by
Linnaeus in the Systema Naturae ( 1758). The Descriptiones . . .were detailed and
extremely competent descriptions of mainly Old World fishes some of which
were well known Scandinavian species. The Genera Piscium and Synonymia nominum
Piscium were, however, the most important as they facilitated the listing of
references to fishes in much earlier literature.
Artedi’s work blazed a trail through the literature and undoubtedly influenced
Linnaeus in his concepts of certain genera and species. Thus, in establishing
Linnaeus’s concept of a given taxon it is frequently necessary to refer to Artedi
(1738). Artedi’s concept of the limit of a genus or species was, however, often
restricted to the earlier literature and has to be assessed within the frame of
contemporary knowledge. It is ignorance of this elementary concept that led
Fricke (1990) to unearth a single early illustration and dependent later
references, in attempting to upset an established usage for Callionymus dracunculus
(Wheeler, 1990), when a secondary type specimen was available in the
Gronovius collection.
This example, however, serves to emphasize an important point when
establishing the limits of Linnaean names, which also has bearing on the
designation of lectotypes when desired. Because the framework of the Systemu
Nuaturae is very largely a compilation, with names of animals based on specimens
examined but mostly on literary sources, sometimes combined, there are a
number of taxa which can be proved to be composite. Where this is established it
is essential that zoologists seeking to restrict the use of the name should be
motivated primarily to stabilize zoological nomenclature, not as in the case cited
to unearth an early illustration cited by Artedi, who was cited in turn by
Linnaeus and invest in it a greater importance than that given to a secondary
source which was referrable to the species for which the Linnaean name was
widely employed.
For workers attempting to establish the present existence and location of type
specimens of fishes named by Linnaeus the following summary may be useful
although details of most surviving specimens are given in Wheeler (1958),
Fernholm & Wheeler ( 1983), Wheeler (1985) and the present paper.
1. References to Amoen. acad., Chin. Lagerstr., and in the case of Muraena hzlena,
Mus. Ad. Fr. refer to specimens that were described in academic dissertations.
156
A. WHEELER
Many of the specimens are still preserved in the Zoological Institute, Uppsala.
2. References by Linnaeus (1758, 1766) to Mus. Ad. Fr. and to Harselqv. itin.
are to specimens of which many are preserved in the Swedish Museum of
Natural History, Stockholm. Most of the 1758 citations are to the first volume of
the Museum Adolph Friderici (Linnaeus, 1754a) but some, which lack page
numbers, refer to the second volume (Linnaeus, 1764a).
3. References by Linnaeus (1 758) to Gron. m u . and (1 766) to Gron. rooph. refer
to specimens described by L. T. Gronovius in the Museum lchthyologium (1754,
1756) and to the <oophylacium Gronovianum (1763). Many of the specimens
described in these works are in the British Museum (Natural History), others are
in the Zoological Museum, Copenhagen (see Wheeler, 1958, 1989).
4.Specimens described and named by Linnaeus (1758), but for which he gave
no literary reference, are in the Zoological Institute, Uppsala, and in the
Linnean Society of London (Swedish species). However, the specimen of Pelecus
cultratus described by Linnaeus (1751) in his Skanska Resi is still preserved in
Uppsala. The majority of the collection of North American fishes made by
Alexander Garden and named by Linnaeus (1 766) is in the Linnean Society of
London.
5. No specimens are known that relate to the descriptions of Artedi ( 1738), the
most frequently cited source for fishes. There is one specimen of Ostracion semu
lato in the British Museum (Natural History) which might possibly have been in
Sloane’s museum when Artedi visited London in 1734 and which he might have
described, but the evidence is tenuous. There is also the possibility that some of
Seba’s specimens examined by Ariedi in Leiden in 1735 might still exist
unrecognized (see Boeseman, 1970), but none have so far been noted. [It might
be noted parenthetically that the only Sloane specimen in the British Museum
(Natural History) which is certainly identified is the holotype of Chauliodus sloanei
(Bloch & Schneider, 1801) which had been illustrated and described by Catesby
( 1743).]
CATALOGUE OF LINNAEAN SPECIMENS IN THE ZOOLOGICAL INSTITUTE, UPPSALA
The specimens listed here include all those in the collection of the Zoological
Institute of the University of Uppsala that have been located and identified as
Linnaean. These specimens were listed by Lonnberg (1896) and by Holm
(1957), but have been re-examined, where possible reidentified and have been
verified through the literature as consistent with being Linnaean material. They
are listed in the order and under the names of higher taxa given by Nelson
(1984), but each taxon is listed within the family by its modern name.
Each species account begins with the modern nomenclature for the taxon,
followed by the relevant references to Linnaeus’s descriptions or references to it,
as well as those significant references by other authors. Details of the specimen
are then given in the following sequence: Zoological Institute, Uppsala (ZIU)
collection number (these follow Holm, 1957 but Lonnberg’s number is given in
brackets), the length of the specimen in mm (SL = standard length; T L = total
length), a note of its condition and state of preservation, and a transcription of
the original label many of which were written by Thunberg. Discussion of the
specimen is restricted mainly to its standing as a type specimen, or the evidence
for it being part of Linnaeus’s collection.
LINNAEAN FISHES I N UPPSALA
157
Petromyzontijormes
Petromyzontidae
LampetraJluuintilis (Linnaeus, 1758)
PetromyzonJluuiatilis Syst. N a t . (10): 230 (1758); Syst. Nat. (12): 394-(1766).
Specimen: ZIU 157 [34]; T.L. 192 mm; in alcohol, poor condition.
Label: early labels missing.
Discussion: This species was named by Linnaeus from the literature and no type specimens would be expected
to exist.
A specimen of Petromyzon Jlulluuiatilis was listed as being in the Alstromer-Linnt donation of 1749 and
subsequent years (Thunberg, 1787) and was included in Thunberg’s manuscript catalogue of the Uppsala
collection of 1828. Lonnberg (1896), however, recorded the specimen as ‘lost’, although it had evidently been
rediscovered by the time Holm (1957) published his catalogue of the zoological collection. While the specimen
may be correctly identified with the Alstromer-LinnC donation it is not a type specimen.
M y x i n i f rmes
Myxinidae
h@xine glutinosa Linnaeus, 1758
Myxineglutinosa Syst. Nat. (10): 650 (1758); Sys. Nat. (12): 1080.
Specimen: ZIU 222 [116]; T.L. 246 mm; in alcohol, poor condition.
Label: Myxine/glutinosa./Mus. Linn.
Discussion: Myxine glutinosa was based on references to the earlier literature including the Museum Adolph2
Fridericiunum. Specimens believed to have originated in the collection of King Adolf Fredrik are still preserved
in the Swedish Natural History Museum and were identified by Fernholm & Wheeler (1983) as having type
status. There seems to be no reason to suspect that the present specimen was part of that collection.
This specimen was listed by Thunberg ( 1 787) as from the combined Alstromer-Linne donation of 1749, in
which he was followed by Lonnberg (1896) and Holm (1957).
Lamnijormes
Scyliorhinidae
Scyliorhinus stellaris (Linnaeus, 1758)
Sgualus catulus Syst. Nat. (10): 235 (1758); Syst. Nut. (12): 399 (1766).
Specimen: ZIU 162 [38]; T.L. 224 mm; in alcohol, good condition.
Label: Squalus/Catulus/Mus. Linn.
Discussion: Sgualus catulus was described from five literary references none of which are associated with the
LJppsala collection. This specimen has no type status.
It is listed as having been part of the Alstromer-Linnt donation of 1749 by Thunberg (1787), by Lonnberg
(1896) who attributed the label on the jar to Thunberg, and by Holm (1957).
Sgualifrmes
Squalidae
Sgualus acanthias Linnaeus, 1758
Sgualus acanthiar Syst. N a t . (10): 233 (1758); Syst. Nat. (12): 397 (1766).
Specimens: ZIU 159 [36]; T.L. 341 mm; in alcohol, moderate condition; ZIU 160 [36]; T.L. 390 mm; dried
stuffed skin very mis-shapen.
Labels: [both specimens] Squalus/acanthias/a/Mus. Linn.
Discussion: Sgualus acanthias was named by Linnaeus from several literary sources including his account in
Myus. Ad. Frid. (Linnaeus, 1754a). A specimen which might be the original of that description was identified by
Fernholm & Wheeler (1983) who consider it to he a putative type specimen.
There is no reason to suspect that either of these two specimens has type status. They originated in the
Ahlstromer-Linnt donation of 1749 (Thunberg, 1787; Lonnberg, 1896 and Holm, 1957).
Specimen: ZIU 56 (see Holm (1957: 58)); T.L. 680 mm; dry, stuffed and mounted.
Label: Squaluslacanthiaslj Mus. Gust. Ad.
Discussion: This specimen was part of the donation of King Gustav IV Adolf, the grandson of Queen Louisa
Ulrika, made when the remainder of the collection of the Queen was removed from Drottningholm. There is no
reason to assume that it has type status, even though other specimens (mainly invertebrates) in that donation
described by Linnaeus (1764b) and now in the Uppsala collection are type specimens.
158
A. WHEELER
Squatinidae
Squafina squatina (Linnaeus, 1758)
Squalus squatina Sysf. Naf. (10): 233 (1758); Syst. Naf. (12): 398 (1766).
Specimen: ZIU 161 [37]; T.L. 228 mm; dry specimen on mount.
Label Squalus/squatina/S/Mus/Linn.
Discussion: Squalus squatina was based on six earlier literary references but none of them were associated with
Uppsala material. This specimen has no type status.
This specimen originated in the Ahlstromer-LinnC donation of 1749 (Thunberg, 1787; Lonnberg, 1896 and
Holm, 1957).
Rajijormes
Rajidae
Raja clauata Linnaeus, 1758
Raja clavata Syst. Naf. (10): 232 (1758); Sysf. &at. (12): 397 (1766).
Specimen: ZIU 158 [35]; T.L. 191 mm; in alcohol, poor condition.
Label: Raja/clavata/Mus. Linn. Two labels in different hands.
Discussion: Raja clavafa was described from several earlier literary accounts none of which were associated with
Uppsala specimens. There is no reason to assume this to be type material.
This specimen was attributed to the Ahlstriimer-Linnt donation of 1749 by Thunberg (1787), Lonnberg
(1896), and Holm (1957). Lonnberg suggested that the specimen might belong to the species Raja radiafa
Donovan. 1806.
Acipmscrifonna
Acipenseridae
Acipenscr sturio Linnaeus, 1758
Acipnscr shrrio Syst. Naf. (10): 237 (1 758); &st. Nat. (12): 403 (1 766).
Specimen: ZIU I70 [48];T.L. 1940 mm; stuffed skin.
Label: Acipenser Sturio/Mus. Linn et Ahlstroem.
Discussion: Aci#m.scr shrrio was described from five literary references, none of which have relevance to the
Uppsala collection. However, two of the references (Linnaeus, 1754a; Gronovius, 1756) relate to specimens
which are still in existence, the former in the Swedish Museum of Natural History (Fernholm & Wheeler,
1983) and the latter in the Gronovius collection in the British Museum (Natural History) (Wheeler, 1958).
Both have claims to be considered to be types. The present specimen is not type material.
This specimen was part of the Ahlstromer-Linnt donation of 1749 (Thunberg, 1787; Holm 1957); it was said
by Liinnberg (1896) to be lost.
ClupelIb7Ynes
Clupeidae
Clujmnodon thrissa (Linnaeus, 1758) [Fig. 31
Clupea fhrissa Syst. Nat. (10): 318 (1758); +sf. Nut. (12): 524 (1766).
Mystus (altus) corpore ovata, Linnaeus (1754b): 26.
Clupea ( Triza) corpore ovato, Linnaeus ( 1759): 25 1.
Specimen: ZIU 107 [30]; S.L. 103 mm, T.L. 115+mm; in alcohol.
Label: Clupea Thrissa/Mus. Lagerstr.
Discussion: According to Thunberg (1787) this specimen was part of the donation of Magnus Lagerstrom of
1748 and was so listed by both Liinnberg (1896) and Holm (1957). However, as discussed earlier this species
was one of the fishes described by Osbeck (1757) and it is my opinion that the present specimen originated in
Osbeck’s collection.
The specimen was first described in the dissertation of Johannes Laurentius Odhelius entitled Chinensia
Lagcrstrhiana (Linnaeus, 1754b). This was reprinted in the Ammifates Academicw (Linnaeus, 1759) but with
the name changed to Clu#ca T k a . As the references cited by Linnaeus (1758) in the formal publication of
C&ea thrissa included the citation of Odhelius’s dissertation in its first printing this specimen must be regarded
as having type status.
Linnaeus (1758) also cited under this binomen the works of Patrick Browne (1756) on the natural history of
Jamaica and Osbeck’s (1757) account of his voyage to China. Because of this geographical range there is no
doubt that the taxon Cl+ thrissa was composite as of its original publication. I t is relevant here to note that
the fint use of the binomen Clupea thrissa to be published was in Osbeck (1757), which could be interpreted as
further evidence that this is an Osbeck specimen.
LINNAEAN FISHES IN UPPSALA
159
Figure 3. Clupunodon thrissu (Linnaeus, 1758) (ZIU 107), type specimen.
Engraulidae
Coiliu mystus (Linnaeus, 1758) [Fig. 41
Clupcu mysfus Syst. Nut. (10):
319 (1758); Syst. Nut. (12): 524 (1766).
Mystus (ensiformis) corpore ensiforme, Linnaeus ( 1754b): 26.
C/upeu (Mystus) corpore ensiforme, Linnaeus (1759): 252.
Specimen: ZIU 108 [31]; S.L. 152 mm, T.L. 163 mm; in alcohol.
Label: Clupea encrasicolus/Mus. Lin.
Discussion: According to Thunberg (1787) a specimen of Clupeu Mysfus was included in the Lagerstrom
donation to the University Museum of 1748. There is a discrepancy between this attribution and the label with
Figure 4. Coiliu mystus (Linnaeus, 1758) (ZIU 108), probably a type specimen.
160
A. WHEELER
the specimen which relates it to the Linnaeus donation. This discrepancy was noted by Liinnberg (1896) who
explained it by suggesting a former confusion between labels and he included the specimen in his list of the
Lagerstriim donation. Holm (1957) likewise attributed this specimen to this source.
In proposing the taxon C~I@Wmystus Linnaeus (1758) cited two earlier published references, Osbeck (1757)
and Odhelius’s dissertation describing the Lagerstrom donation (Linnaeus, 1754b); he also included a line of
meristic data without refaence to a published description which I interpret (Wheeler, 1985) as showing that he
had examined a specimen himself. In this case the meristic data are identical, with one minor exception, with
those given in Linnaeus ( 1754b).
Earlier, I have suggested that the specimens described for Odhelius’s dissertation were those collected by
Osbeck and given to Lagerstrijm. If this is accepted then there would have been only one specimen for both
sources, and Linnaeuo’si meristic data might well refer to a -nation
of that specimen. In these
circumstances I fed that the pnsent specimen can be regarded as probably the single reference specimen for
specimen of Mtlr&a h e l m Linnaeus, 1758.
0
I.
LINNAEAN FISHES I N UPPSALA
161
Osbeck’s (1757) and Odhelius’s (Linnaeus, 1754b) descriptions. T h e only qualifications to be made in
regarding this specimen as the type of the species, are the confusion over the source of the specimen (noted
above) and the presence of a second later acquired specimen in the University Museum. This second specimen
of Clupea m p t u s was included in Thunberg’s donation of 1775 (Thunberg, 1787: 32). T h a t there were two
specimens is confirmed in the systematically arranged manuscript catalogue of the collection compiled by
Thunberg in 1828 where two specimens of Clupea Mystus are listed, a from Lagerstrom, B from Thunberg. This
casts an element of doubt over recognizing the present specimen as type of the species but the probability seems
to be in favour of regarding it as such.
Anguilliformes
Muraenidae
Echidna sp. [Fig. 51
Muraena helena (part) Syst. N a t . (lo): 244 (1758); Sysf. N d . (12): 425 (1766).
Muraena pinnis pectoralibus carens, Linnaeus (1746): 4 0 (1749): 319.
Specimen: Z I U 57 [53]: S.L. 322 mm, T.L. 324; mm; in alcohol.
Label: Muraena/Helena./Mus. Ad. Frid.
Discussion: In naming the species Muraena helena Linnaeus (1758) cited five published references. T h e second of
these was given as Mus. A d . Fr 1. p. 319. Although most references in this form refer to the folio Museum Adolphi
Friderici (Linnaeus, 1754a) the pagination given clearly refers it to Laurentius Balk’s dissertation reprinted in
the Amoenitates academicae (Linnaeus, 1749a), which is often otherwise referred to as “Museum Principis”. This
dissertation referred to the collection of specimens donated by Adolf Fredrik in 1745. There is thus reason to
expect the specimen of Murama to be in the collection of the Zoological Institute.
A specimen labelled Murama Helena was listed in the 1745 donation of the Crown Prince by Thunberg (1787)
and also in the manuscript catalogue compiled by Thunberg in 1828. Lonnberg (1896) also noted the specimen
but recognized that it was not conspecific with M u r a m heleno of authors, and suggested that it was identifiable
as MuraenapolyZona Richardson, 1845. Holm (1957) also listed the specimen as from the donation of 1745.
There seems to be no doubt that this specimen is a type, and indeed is the only surviving type specimen of
Muraena h e h a . Equally, there is no doubt that it is referable to the genus Echidna, on account of its dentition,
with rounded blunt teeth in both the jaws and on the vomer.
Salmonlfonnes
Salmonidae
Salvelinus alpinus (Linnaeus, 1758)
Specimen: ZIU 206 [102]; S.L. 221 mm, T.L. 259 mm; in alcohol.
Label: Salmo/lacustris/Mus. Linn.
Discussion: This specimen was presumably the one included in Thunberg’s (1787) listing of the
Alstromer-Linni: donation of 1749 under the name ofSalmo lacustris. I t was similarly listed by Lonnberg (1896)
and Holm (1957). However, it is not a type specimen of Salmo lacustris Linnaeus, 1758 which was based entirely
on Artedi’s description. Presumably the label was written by Thunberg while curating the collection and this
was his identification.
Osmeridae
Osmerus eperlanus (Linnaeus, 1758)
Salmo eperlanus $st. N Q ~(10):
.
310 (1758); Syst. N a t . (12): 511 (1766).
Specimen: Z I U 207 [103]; S.L. 166 mm, T.L. 196 mm; in alcohol.
Label: Salmo eperlanus Mus. Linn. a
Discussion: Salmo eperlantcs was based on two literary references neither of which were associated with the
Uppsala collection. No type specimen is to be expected in this material.
This specimen is part of the Ahlstromer-Linne donation of 1749 according to Thunberg (1787), Lonnberg
(1896), Holm (1957) and Thunberg’s manuscript catalogue of 1828.
Charactfonnes
Erythrinidae
Hoplerythrinus unitaeniatus (Spix, 1829)
Cyprinus c e p h a h , part. syst. N Q ~(10):
.
322 (1758); $st. Naf. (12): 527 (1766).
Specimen: ZIU 213; S.L. 162 mm; T.L. 195 mm; in alcohol.
Label: Indecipherable; no external label.
Discussion: This specimen is listed by Holm (1957) as being from the 1749 donation of Ahlstromer-Linnt
material, but if listed it is not readily identifiable in the earlier lists of Thunberg (1787) and Lonnberg (1896).
162
A. WHEELER
This specimen has already been dixussed by Fernholm & Wheeler (1983) in the context that another
specimen of H.plnstAr;nur unitutn&us is in the collection of the Swedish Natural History Museum, labelled
CyPriu caplurluc, and is certainly a type specimen of that taxon. I t is possible that this Uppsala specimen had a
common origin with the Stockholm specimen and that it also is a type specimen of Cyprinus cephulus, but as its
curatorial history is incomplete this cannot be established with certainty.
Gymnotifoms
Gymnotidae
Gymnotus curupo Linnaeus, 1758 [Fig. 61
Gymnotuscurufi Syst. Nut. (10): 246 (1758): Syst. Nut. (12): 427 (1766).
Specimen: ZIU 56 [52]; S.L. 295 mm, T.L. 298 mm; in alcohol.
Label: Old label missing.
Discussion: According to Holm (1957) and Lonnberg (1896) this specimen originated in the donation of the
Adolf Fredrik in 1746. Thunberg (1787), however, did not list it in that donation assuming that is was the
specimen of “Gjmnotus electrim” from the Museum Grillianum (Linnaeus, 1748a). As Lonnberg pointed out the
Museum Grillianum specimen was much larger than the present specimen, and the illustration published by
Linnaeus (1749a) from the donation of 1746 fits this specimen exactly.
GynuroruF curupo was based on three earlier printed descriptions one of which was to Amoenitules Acudemicac ( 1,
p. 318, t. 14, fig. 6) which was the reprinted version (Linnaeus, 1749a) of Balk‘s dissertation Museum
Adolpho-Fridn;caRunr (Linnaeus, 1746a). This present specimen appears to be this specimen and it must be
regarded as having type status. I t possesses greater importance because the other surviving specimen with type
status (Swedish Natural History Museum NRM LP 64)is referrable to Eignnaniu viridescms Valenciennes,
1847 (see Fernholm & Wheeler, 1983). Fernholm & Wheeler discussed the Uppsala specimen and concluded
that it had to be regarded as the type specimen of Gymnotus curu#o Linnaeus, 1758.
Cfiriniformes
Cyprinidae
Curarsius uurutus (Linnaeus, 1758)
Cyprinus uurutus Syst. Nut. (10): 322 (1758); Syst. Nui. (12): 527 (1766).
Specimen: ZIU 109 [32]; S.L. 136 mm, T.L. 170 mm; in alcohol.
Label: Internal modern label Curarsius uurutus? (L.). Mus.Linn.
Discussion: This specimen is listed by Holm (1957) as having originated in the donation of Lagerstrom in 1754.
Lonnberg (1896) claimed that the specimen was missing at that date, but it was one of three lots (lot B) listed in
the manuscript catalogue compiled by Thunberg in 1828.
The specimen was referred to in Odhelius’s dissertation C h i m a Lognshiimiunu (Linnaeus, 1754b) as Cyprinus
uureus and in the reprinted edition in Amoenitates Acudemicac (Linnaeus, 1759) as C. uurutus. However, the
dissertation was not cited by Linnaeus in establishing the name Cfirim uurutus in 1758, and the specimen
therefore has no type status.
Specimens: ZIU 60 [57]; S.L. 77, 71, 53, 49, 42 mm, T.L. 101, 91, 71, 74, 57 mm; in alcohol.
Label: Cyprinuslauratus a/Mus. Ad. Frid.
Discussion: These five specimens are attributed to the donation of Adolf Fredrik in 1746 by Holm (1957) as
they are by Lonnberg (1896) and Thunberg (1 787). They were mentioned with a short diagnosis in Balk‘s
dissertation Museum Adolpho-Fredericiunum (Linnaeus, 1746a) and the reprinted edition in Amoenitates Acudemicae
(Linnaeus, 1749a). However, Balk’s dissertation was not one of the published references cited by Linnaeus
(1758) in the naming of Cyprinus uuratus. These specimens therefore have not type status.
These goldfish specimens all have elaborate triple caudal fin lobes, one dorsal and two ventral lobes; one of
them has a double anal fin, and the largest one has a very short based dorsal fin. Evidently, these early
specimens in northern Europe were already subject to considerable cultured variation.
P e l e m cultrutus (Linnaeus, 1758)
Cyprinus cultrutus S y s . Nut. (10): 326 (1758); Syst. Nut. (12): 531 (1766).
Specimen: ZIU 224; S.L. 206 mm, T.L. 235 mm; in alcohol, poor condition.
Label: Cyprinus cultratus.
Discussion: The name Cyprinus cultrutus was based by Linnaeus on his own earlier description in S k h k u
Resu.. . (Linnaeus, 1751). This description was of a fish given to Linnaeus during his journey in SkHne by
borgmbtre Schulten i Kristianstd which was captured at the river Helgeh in 1749 (the year Linnaeus was in
SkHne). The specimen was illustrated (pl 2) in the Skcfnsku Resu . ..There is no doubt that this is the original
specimen which was listed by both Liinnberg (1896: 45) and Holm (1957), even though Lonnberg noted that it
had been lost and refound several times. I t is therefore the holotype of the species Cyprinus cultrutus.
LINNAEAN FISHES IN UPPSALA
163
Figure 6. Gymnotus camp0 Linnaeus, 1758 (ZIU 56), type specimen.
Phoxinus phoxinus (Linnaeus, 1758)
Cyjrinus aphya Syd. Nat. (10): 323 ( 1 758): Syst. N a t . (12): 528 ( 1 766).
Specimen: ZIU 2 11 [ 1121; S.L. 7 1 , 62 mm, T.L. 79 , 74 mm; in alcohol; poor condition.
Label: Cyprinus/Aphya/Mus. Linn.
Discussion: These specimens are in bad condition and have evidently been dry at some time. Their condition is
such that it is not possible to be certain of their identity although they are tentatively identified with Phoxinus
phoxinus because Cyprinus ap/ya is usually regarded as a synonym of that taxon. However, the scales which are
visible seem rather large for P. phoxinw of this size.
These specimens were listed by Holm (1957), Lonnberg (1896), and Thunberg (1787) as originating in the
Alstromer-Linni donation of 1749. There is no evidence that they are type specimens of either Cypinusphoxinus
or C. aphya as neither taxon was founded on literary references to the Linnaean dissertations which were the
foundation of much of the Uppsala collection.
+
Lcuciscine indet.
Specimen: ZIU 212 [115]; S.L. 204 mm, T.L. 235 mm; in alcohol, fair condition.
Label: Cyprinus cephalus.
Discussion: This specimen was listed by Lonnberg (1896) and Holm (1957) as having been part of the
Alstromer-Linni: donation of 1749. Lonnberg was, however, unsure of its identification and suggested that it
might have been a hybrid, adding that as its original label was lost it is not certainly a type specimen. There is
no doubt that it is not a type specimen of Cyprhus cephalus, for none of the literary references cited for that taxon
refer to material which would be in Uppsala.
For discussion of other type specimens of C. cephalus see Fernholm & Wheeler (1983: 212) and the present
paper under Hoplerythrinus unitaeniatus (p. 161).
Cobitidae
Cobitis taenia Linnaeus, 1758
Cobitis taenia Syst. Nat. (10): 303 (1758); Syst. Naf. (12): 499 (1766).
Specimen: ZIU 205; S.L. 76 mm, T.L. 84 mm; in alcohol, fair condition.
Label: Cobitis/taenia/Mus./Linn.
Discussion: This specimen was listed by Holm (1957), Lonnberg (1896) and in Thunberg’s manuscript
catalogue of 1828, as having originated in the Alstromer-Linne donation of 1749.
The name of Cobitis taenia was based on three references to earlier literature, Artedi (1738), Linnaeus (1746b)
and Gronovius ( 1 754). In addition there is a set of meristic data which suggests that Linnaeus had examined a
specimen himself. As no specimen of Cobitis tamia is preserved in Linnaeus’s personal collection (Wheeler, 1985)
I64
A. WHEELER
it is possible that this specimen was the one used by Linnaeus to derive these data either before or after the
donation of 1749. There is no way ofestablishing that it was a Linnaean specimen, but the evidence in favour
of this is strong. I t may be a type specimen.
Siluriifomes
Aspredinidae
Aspredo aspredo (Linnaeus, 1758) [Fig. 71
Silurus aspredo Syst. flat. (10): 304 (1758); Syst. Nai. (12): 502 (1766).
Specimen: ZIU 48 [43]; S.L. 226 mm, T.L. 242 mm; in alcohol, good condition.
Label (not original); Silurus asprcdo.
Discussion: Silurus aspedo was named by Linnaeus (1758) on three earlier descriptions, Gronovius (1754), Klein
(1749), and to the reprinted edition of Balk’s dissertation Museum Adolph-Fridmicianum in Amoenitates Academicae
(Linnaeus, 1749a).
The present specimen was listed by Xolm (1957), Lonnberg (1896) and Thunberg (1787) as having
originated in the 1745 donation of the Crown Prince Adolf Fredrik. I t was therefore one of the specimenswhich
formed the basis of Laurens Balk’s dissertation (Linnaeus, 1746a) and is clearly the figured specimen in this and
the later edition (Linnaeus, 1749a). I t therefore must be regarded as having type status, and as it is the only
specimen surviving from the three cited works has to be regarded as the type.
A second specimen of the species which probably came from the same source is preserved in the Swedish
Natural History Museum, Stockholm (Fernholm & Wheeler, 1983) but this is not a type specimen. It seems
probable that there were two specimens in the Crown Prince’s collection before 1745, one of which he donated
to Uppsala University (which later became this type specimen) while he retained the other which eventually
passed to the Stockholm collection.
Callichthyidae
Hoplostemum littorale (Hancock, 1828)
Specimen: ZIU 55 [51]; S.L. 66 mm, T.L. 88 mm; in alcohol, fair condition.
Label in Thunberg autograph: Silurus callichthys Mus. Thumb.
Discussion: This specimen is listed by Holm (1957) as a specimen from the donation of Adolf Fredrik, when
Crown Prince, in 1745. A similar, but very confused statement, by Lijnnberg (1896) probably caused Holm to
accept this fish as belonging to the 1745 donation. Lonnberg wrote under the heading Callichthys “the orginal
specimen from Museum PrinCipu must either have been lost very early as it is not recorded in Thunberg’s
catalogue.. .,or there can be another possibility as there is a specimen of Callichthys in the “old collection”
of the Uppsala Museum. But this is labelled by Thunberg himself, as belonging to “Museum
LINNAEAN FISHES IN UPPSALA
165
Thunbergianum” . . . It is very possible that this is the specimen from Museum Principis as it is very similar to the
figure Tab. XIV, fig. 1”. In this, Lonnberg appears to be arguing that although the original 1745 donation
specimen may have been lost (as it was not catalogued by Thunberg) and this specimen appears to have been
donated by Thunberg (in 1775), it is very possible that it is nevertheless the original ( 1 745) specimen.
However, the only evidence is against this supposition. Firstly, no specimen of Silurus callichthys is listed in
Thunberg’s (1787) listing of the Adolf Fredrik donation, or in his manuscript catalogue of 1828. Secondly,
Thunberg (1787) lists a specimen ofSilurus cakchtus in his own donation of 1775. Thirdly, the bottle is labelled
as from the Museum Thunbergianum in Thunberg’s autograph. Fourthly, despite Lonnberg’s comment on the
similarity of the specimen to the figure in Linnaeus (1749a), there are considerable discrepancies, notably in
the length of the barbels, the size of the eye and the curvature of the back. It is my opinion that this is not the
specimen described by Linnaeus (1749a), which appears to have been referrable to the genus Callichthys
whereas this specimen is Hoplosternum littorule.
The name Silurus ca&hthys was based by Linnaeus (1758) on three earlier references of which Balk’s
dissertation Museum Adolpho-Fridericianum (Linnaeus, 1749a) was first named. The others were to Gronovius
(1754) and Barrert (1741). The Gronovius specimen is still present in the British Museum (Natural History)
(1853.11.12.194); see Wheeler (1989). As the specimen described in Balk’s dissertation is no longer in the
Uppsala collection this Gronovius specimen appears to be the only surviving type material.
GadtJomes
Gadidae
Lota lota (Linnaeus, 1758)
Gadus h a Gst. Nat. (10):255 (1758); *st. Naf. (12): 440 (1766).
Specimen: ZIU 178 [61]; T.L. c 120 mm; in alcohol (this fish hangs in a sealed bottle which has not been
opened since the eighteenth century).
Discussion: According to Holm (1957), Lonnberg (1896) and Thunberg (1787) this specimen belongs to the
Alstromer-Linnt donation of 1749.
The name Gadus lota was founded on three citations to earlier literature, a description in Fauna Suecica
(Linnaeus, 1746b), and to two parts ofArtedi’s Zchthyologiu (1738). Wheeler (1985) considered that it was likely
that the dry skin of a specimen in Linnaeus’s personal collection was probably the specimen described in Fauna
Suecica. However, there is no proof in support of this as Linnaeus’s specimens were inadequately labelled, and it
is possible that the present specimen is as likely to have been referred to in that book. It is marginally less likely
to be a type specimen of this common Swedish fish than the specimen which was retained in Linnaeus’s
personal collection.
Merlangius merlungus (Linnaeus, 1758)
Gadus merlangus Gst. Nat. (10); 253 (1 758); syst. Nut. (12): 438 (1 766).
Specimen: ZIU 175 [56]; S.L. 205 mm, T.L. 232 mm; in alcohol, poor condition, head partly missing.
Label: Gadus/merlangus/Mus. Linn.
Discussion: This specimen is part of the 1749 donation of Alstromer-Linnt according to Holm (1957),
Lonnberg (1896) and Thunberg ( 1 787).
The name Gadus merlangur was based on five earlier publications, Artedi (1738), Gronovius (1754) and
Linnaeus (1746b, 1747, 1751). Two specimens of this species are preserved in the Gronovius collection
(Wheeler, 1958) which may have type standing. The three Linnaean references, two of which (Linnaeus, 1747,
1751) are notes made on travels in Sweden, might refer to this specimen, and it is possible that it is part of the
type series. However, in view of its condition it seems undesirable to strive to prove that this specimen has type
standing.
Merlangius merlangus (Linnaeus, 1758)
Specimen: ZIU 176 [59]; S.L. 188 mm, T.L. 205 mm; in alcohol, in bad condition.
Labels: GaduslcarbonariuslMus. Linn, at head and foot of jar.
Discussion: This specimen is part of the 1749 donation of Alstromer-Linnt according to Holm (1957) and
earlier authors. It has no type status, as discussed above for Gadur mcrlangus, and G. curbonurius was based solely
on Artedi’s (1738) generalized account. G. carbonarius is usually regarded as a synonym of G. uirens (Pollachius
uirens) .
This specimen is in very bad condition. Lonnberg (1896) suggested that it was probably referrable to Gadus
merlangus, a view with which I concur. However, the only distinctive features which are discernable are the long
lower jaw and the anal fin origin which is opposite the middle of the first dorsal fin.
Pollachius pollochius (Linnaeus, 1758)
Gadus pollachius Gst. Nat. (10): 254 (1758); Syst. Nat. (12): 439 (1766).
Specimen: ZIU 177 [60];S.L. 150 mm, T.L. 165 mm; in alcohol, in good condition
Labels: Gadus/Pollachius/Mus.Linn, plus second label indecipherable.
166
A. WHEELER
Discussion: This specimen is part of the 1749 donation of Alstromer-Linnt according to Holm (1957) and
earlier authors.
The name Gadus polfachius was based on three earlier references, Artedi (1738), Gronovius (1754) and
Linnaeus (1747). The last of these was the account of Linnaeus’s journey to West Gothland (Wiirg6fa-Resa.. .)
and it is possible that this specimen was collected on that journey and presented to the museum two years later.
It may therefore have type status.
Raniceps raninus (Linnaeus, 1758)
Blmnius raninus Sysf. Nut. (10): 258 (1758); Syst. Nat. (12): 444 (1766).
Specimen: ZIU 181 [MI; S.L. 259 mm, T.L. 283 mm; in alcohol, in moderate condition.
Labels: Blennius/raninus/Mus. Linn. (internal label “Raniceps raninus (L) = Blennius raninus L =
Batrachocephalus blennoides Mus. Linn”.)
Discussion: Blmnius raninus was described and named in 1758 with no literary references but the locality was
given as “Habitat in Sveciae lacubus”. The description was detailed and clearly Linnaeus had a specimen to
examine. There is a specimen of this species in Linnaeus’s personal collection (Wheeler, 1985) but because it
was not labelled and has no Linnaean annotation he decided it should not be regarded as a type specimen. The
Uppsala specimen is very probably the type, however, as it forms part of the Alstromer-Linnt donation of 1749
(Holm, 1957) and was thus available to Linnaeus long before the publication of the S y s t m Nafurae (1758).
Lophiifonnes
Antennariidae
Histrio histrio (Linnaeus, 1758) [Fig. 81
Lophius histrio Sysf. Nut. (10): 237 (1758): Sysf.N a f . (12): 403 (1766).
Specimens: ZIU 100 [21]; S.L. 74, 26, 18 mm, T.L. 109, 36, 24 mm; in alcohol, in moderate condition except
for the smallest which is poor.
Labels: Lophius/histrio/Mus. Lagerstr.
Discussion: The name Lophius hishio was based by Linnaeus on six literary references. Of those that are relevant
to Swedish collections two might be considered here (the remainder having been discussed by Fernholm &
Wheeler, 1984). The first is to Linnaeus (1754b) which was Odhelius’s dissertation on the Lagerstrom
collection, and the second to Osbeck‘s ( 1 757) Dagbok ojiver m Osfindisk-Resa. . . .
These specimens are labelled as kom Lagerstrom’s collection and are listed as such by Holm (1957),
Lonnberg (1896) and there is an entry under Lophius histrio a attributed to Lagerstrom in Thunberg’s ms
catalogue of 1828, but it is not certain that this indicates a single specimen or one lot of specimens. Thunberg’s
1828 catalogue also contains an entry for B from Thunberg’s collection. I suspect, in view of this, and the
differing states of preservation of the specimens in this jar that at some time the a and B lots may have been
aggregated. This introduces a complication into the recognition of type material because there is little reason to
Figure 8. Histrio histrio (Linnaeus, 1758) (ZIU IOO), one, possibly two of these specimens are types.
LINNAEAN FISHES I N UPPSALA
167
doubt that the original Lagerstrom specimen(s) is the material described in Odhelius’s dissertation and
therefore has type status (as Lonnberg (1896: 30) has already pointed out).
This is another example in which material at Uppsala is listed by both Osbeck (1757) and in the Lagerstrom
donation (Linnaeus, 175413).As discussed earlier it is my considered opinion that both references referred to the
same specimen which was collected by Osbeck.
The identification given of these specimens is tentative.
Ogcocephalidae
Ogcocephalus uespertilio (Linnaeus, 1758)
Lophius uespertiho Syst. Nat. ( 10): 236 ( 1 758); $st. Nat. ( 12) : 402 ( 1766).
Specimen: ZIU 163 [40]; S.L. 184 mm, T.L. 215 mm; dry, mounted on wooden block (see Fig. 2).
Labels: Lophius/Vespertilio/Mus. Linn. (in duplicate).
Discussion: The name Lophius uespertilio was based by Linnaeus on six literary references of which only that to
Linnaeus (1754a), the Museum Adolphi Friderici volume I is at all relevant to Swedish collections. Material from
the collection of Adolf Fredrik described in this work is, however, in the Swedish Museum of Natural History
(Fernholm & Wheeler, 1984) and there are no known examples from this collection to be found in Uppsala. It
serms unlikely therefore that this specimen is involved in the typification of Lophius uespertilio.
It was listed as having originated in the donation known as the Alstromer-LinnC collection which contains
few (if any) type specimens of Linnaeus’s species (see Holm, 1957) and neither Holm nor Lonnberg (1896)
claim it to be a type specimen. Despite this, and perhaps because of a misunderstanding, Bradbury (1967)
claims that this specimen is the holotype of the species. It cannot be the holotype, and is very unlikely to be a
type specimen of any kind.
Cyprinodontijomes
Exocoetidae
Exocoetus volitans Linnaeus, 1758 [Fig. 91
Exocoetus uolitans $st. Nat. (10): 316 (1758); Syst. &at. (12): 520 (1766).
Specimen: ZIU 59[55]; S.L. 174 mm, T.L. 186+mm (tail broken); in alcohol.
Label: No old label.
Discussion: The name Exocoetus uolitanr was based by Linnaeus on three earlier literary references, the
Amoenitates Academicae reprint of Balk’s dissertation (Linnaeus, 1749a), Artedi (1 738) and Gronovius (1 754).
According to Lonnberg (1896) and Holm (1957) the present specimen originated in the donation of the Crown
Prince Adolf Fredrik in 1745 and was therefore the specimen described in Balk‘s dissertation (Linnaeus, 1746a).
It therefore has type-standing.
Figure 9. Exocoetus uolitans Linnaeus, 1758 (ZIU 59), type specimen.
168
A. WHEELER
Bruun (1935) has discussed this specimen (as did Lilljeborg (1891) previously), in the light of the composite
nature of Linnaeus’s Exocoetus volifonr (he regarded the meristic details given by Artedi as evidence of a second
species). I t appears therefore that this specimen should be regarded as the type of the species.
Belonidae
Belone belone (Linnaeus, 1758)
Esox belloneSyst. Nut. (10): 314 (1758); Syst. Nut. (12): 517 (1766).
Specimen: ZIU 208 [104]; S.L. 323 mm, T.L. 345 mm; in alcohol.
Labels: Esox/belone. a/Mus. Linn.
Discussion: Esox bellone (corrected to E. belone by Linnaeus (1766)) was based on references to Artedi (1738),
Linnaeus (1746b), and Gronovius (1754). There is no reason to suppose that this fish is the one referred to by
Linnaeus (1746b) in Fauna Svccica and as it has no relevance to the other cited literature, there is no cause to
suspect that it is a type specimen. However, Holm (1957), Lonnberg (1896), Thunberg’s ms Catalogue of 1828,
and Thunberg (1787) all attribute this specimen to the Alstriimer-LinnC donation of 1749 and later years. The
label, with its attribution to Esox belone a suggests that it was labelled following Bloch & Schneider (1801) who
recognized two varieties (a and j)of E. belone; it may therefore date only from Thunberg’s curatorship.
Lonnberg (1896) considered that this specimen was not referrable to Ra@histmna belone ( = Belone belone) and
tentatively aligned it with R. truncatu (Lesueur, 1821).
Atheriniformes
Atherinidae
Athnina sp.
Specimen: ZIU 209 11051; S.L. 107 mm, T.L. 118 mm; in alcohol, has been dry.
Label: Atherina/hepsetus/Mus. Linn.
Discussion: As noted above this bottle is labelled Atherina hepsetus, and it is listed under that name by both
Lonnberg (1896) and Holm (1957). That taxon was based on references to Artedi (1738), Gronovius (1754),
and Hasselquist (1757), and to a set of meristic features not clearly attributed to any source. Specimens
representing A. hpsetus are present in the Swedish Museum of Natural History (Fernholm & Wheeler, 1984),
and a specimen of Athmna presbyter (believed to be part of the type series of A. hepsetus) is present in the
Gronovius collection (Wheeler, 1958). The Uppsala specimen cannot be proven to have type status, although it
might have been the source of the meristic data given by Linnaeus (1758). It is not referrable to A . hepsetus
Linnaeus s m u strict0 as the ascending pre-maxillary process is short and broad, and may be identifiable with
A. presbyter Cuvier in C . & V.,1829. In these circumstances there seem to be little point in trying to establish its
type status.
Both Lonnberg (1896) and Holm (1957), as well as Thunberg’s manuscript catalogue of 1828, all list this
specimen as having been in the Alstromer-Linnk collection. There is therefore a probability that it was the
specimen examined by Linnacus to obtain the meristic data given in 1758.
Syngnathgonnes
Syngnathidae
Sygnathus DGUS Linnaeus, 1758
Specimen: ZIU 169 [47]; T.L. 233+mm (section of tail missing); dried specimen (8).
Label: Syngnathur/pelagicusrp/Mus.Linn.
Discussion: This specimen is referrable to the European pipefish S’gnurluLs
wu, independently confirmed by
the author, but alao listed as such by Lonnberg (1896). Despite its labd it is not Syngnsthuspslagicus Linnaeus,
1758, a species which was named from Osbeck’s (1757) description. The pnsent specimen has 33 dorsal fin rays
(25-34 range, but 28-31 in 86% of specimens in S. m,[Dawson, 19821) and at 233 +mm T.L. is considerably
longer than the maximum given for S.pclugicus by Dawson (1982).
This specimen is attributed to the Ahlstromer-LinnC donation of 1749 and later, by Holm (1957) and
Lonnberg (1896).
It is neither the type of S. &CILT nor S. pelagicus.
Syngnathus bphle Linnaeus, 1758
Specimen: ZIU 168 [%I; head length 41 mm; specimen otherwise dried and fragmented.
Label: Syngnathuslacus a/Mus. Linn.
Discussion: Both the names Spgwthvr t@hle and S.acus were based by LInnaeus ( 1 758) on references to Artedi
( 1738) and to Gronovius ( I 754) pad in neither case is there any reason to p u m e that type material would be
in the Uppsala collection. In view of the misidentification of the specimen it can only be assumed that it is a
later addition to the collection and it is doubtful that it is really a specimen from Linnaeus’s collection. It was,
LINNAEAN FISHES IN UPPSALA
169
however, listed by Thunberg in his manuscript catalogue of 1828 as from the Ahlstromer-Linne donation (this
list distinguishes between specimens alpha and beta, and is presumed to be the origin of the a on the label.
It was listed from this donation by Lonnberg (1896) and Holm (1957).
Gusterosteiyormes
Gasterosteidae
Gasterosteus aculeatus Linnaeus, 1758
Gacterosteus uculeutus Syst. Nut. (10): 295 ( 1 758); Syst. Nut. ( 12): 489 ( 1 766).
Specimen: ZIU 201 [92]; five specimens S.L. 51, 51, 56, 54, 56 mm, T,L. 58, 58, 64, 63, 67; in alcohol in
moderate condition.
Label: Gastrosteus/aculeatus/Mus. Linn.
Discussion: The name Gusterosteus aculeatus was based on two earlier references Artedi ( 1 738) and Linnaeus’s
Fauna Suecicu (1 746b). Wheeler (1985) indicated that the three skins in the Linnean Society of London, being
Linnaeus’s personal collection, should be regarded as syntypes of this species.
The present specimens are part of the Ahlstromer-Linni donation of 1749 (Lonnberg, 1896; Holm, 1957;
and Thunberg, 1787) but there is no reason to assume that they are type material.
Scorpaenijormes
Scorpaenidae
Scorpaena scrofa Linnaeus, 1758
Specimen: ZIU 199 [89]; S.L. 122 mm, T.L. 156 mm; in alcohol.
Label: Perca/cottoides/Mus. Linn., modern label Scorpaena scrofa L. = Perca cottoides Mus. Linn.
Discussion: This specimen is listed as part of the Ahlstromer-Linne donation of 1749 and later, by Thunberg
(1787) and later authors. It is unlikely to be the type specimen of Percu cottoides Linnaeus, 1758, which was
based on a specimen in the collection of King Adolf Fredrik (see Linnaeus, 1764a) which if it existed would be
in the collection in the Swedish Natural History Museum, Stockholm, although it is not listed by Fernholm &
Wheeler (1983). Probably the identfication as Perca cottoides was a later addition by Thunberg.
It is not the type specimen of Scorpuenu scrofu Linnaeus, 1758, which name was based on earlier descriptions
by Gronovius ( 1754) and Artedi (1738).
Triglidae
Peristedion cutaphractum (Linnaeus, 1758)
Triglu cutuphructa Sjst. Nut. (10): 300 (1758); Sjst. Nat. (12): 496 (1766).
Specimen: ZIU 204 [95]; S.L. 167 mm, T.L. 181 mm; dry, mounted specimen.
Label: Trigla/cataphracta/Mus. Linnaeus.
Discussion: Linnaeus’s Trigla cutuphracta was based on previous descriptions by Artedi ( 1738) and Gronovius
(1754), and probably mostly on Linnaeus’s then unpublished account which appeared in Linnaeus (1764a).
This last account was based on a specimen which is now in the Swedish Museum of Natural History (Fernholm
& Wheeler, 1983). There is no reason to look for a Linnaean specimen of this species in Uppsala and this
specimen has no standing as a type.
Although listed by Holm (1957), Lonnberg (1896) did not list this specimen but indicated that No. 95
(which number was referred to this specimen by Holm) was lost. A specimen of Triglu cutuphructa was listed
from the Ahlstromer-Linni donation of 1749 by Thunberg ( 1787).
+
Cottidae
Myoxocephulus scorpius (Linnaeus, 1758)
Cottus scorpius Syst. Nut. (10): 265 (1758); Syst. Nut. (12): 452 (1766).
Specimen: ZIU 188 [74]; T.L. c. 200 mm; in spirit, very poor condition in tightly fitting bottle and not
removed.
Label: Cottus/scorpius/a/Mus.Linn.
Discussion: This specimen was listed by Holm (1957) and Lonnberg (1896) as from the Ahlstromer-Linne
donation of 1749, and a specimen of the same taxon was listed by Thunberg (1 787) from this donation. There
is little doubt that this specimen originated from this source. However, there is no reason to suppose that it is a
type specimen as this name was based on four earlier literary references none of them works associated with the
Uppsala collections. Specimens which have claim to type status exist in the Stockholm collection (see Fernholm
& Wheeler, 1983) and also in the Gronovius collection (Wheeler, 1958).
170
A. WHEELER
Myoxocephalus quudricomis (Linnaeus, 1758)
Cottus quadriconriS Sys. Nut. (10): 264 (1758); Sysf. Nut. (12): 451 (1766).
Specimens: ZIU 187 [73]; S.L. 133 mm, T.L. 156 mm; in spirit, moderate condition.
Labels: Unreadable.
(Alsotwo specimens of Cullzonymus l y u ; S.L. 39,40 mm, T.L. 48, 50 mm. These are referred to by Holm (1957)
who seemed to assume they were conspecific.)
Discussion: Linnaeus’s Cottus quadricomis was based on four earlier published accounts (Linnaeus, 1746b, 1748b
and 1754a; and Artedi, 1738). None of these relate to specimens likely to be found in the Uppsala collection,
although one (Linnaeus, 1754a) concerned specimens in the Stockholm collection which are still preserved
there (see Fermholm & Wheeler, 1983). There is no reason therefore to suggest that this specimen has type
status.
According to Holm (1957), Lonnberg (1896) and Thunberg (1787) a specimen of Cottus quudiicomis was
present in the Ahlstromer-Linnt donation of 1749.
Agonidae
Agonus curOgnructus (Linnaeus, 1758)
Cottus cutuphructus Syst. Nut. (10): 264 (1758); Syst. Nut. (12): 451 (1766).
Specimen: ZIU 186 [72]; S.L. 141 mm, T.L. 141+mm; dried whole specimen.
Discussion: The name Cottus cutuphructus was based on two earlier references, Artedi ( 1 738) and Linnaeus
( 1 754a), neither of which concerned specimens which might have been expected to be in Uppsala. Fernholm &
Wheeler (1983) showed that specimens in the Stockholm collection include a type specimen from Linnaeus’s
(1754a) description. Wheeler (1985) discussed a specimen in Linnaeus’s personal collection which is not
thought to have type status.
Holm (1957) liited this specimen as part of the Ahlstromer-Linnt donation of 1749, and a specimen of this
taxon was liited from the donation by Thunberg. Lonnberg (1896) failed to find this specimen and listed it as
lost.
Cyclopteridae
Cyloptms lumpus Linnaeus, 1758
Cycfloptmcs lumpus Qst. Nut. (10): 260 (1758); Syst. Nut. (12): 414 (1766).
Specimen: ZIU 167 [45]; S.L. 138 mm, T.L. 162 mm; dry whole specimen mounted.
Label: Cyclopterus/lumpus./Mus. Linn.
Discussion: This specimen is part of the Ahlstromer-Linnt donation of 1749 (Holm, 1957; Lonnberg, 1896); a
specimen of this taxon being listed from the donation by Thunberg (1787).
This species was named from citations to five earlier works (Linnaeus, 1746b, 1751 and 1754a; Artedi, 1738;
and Gronovius, 1754). None of these relate to specimens expected to be in the Uppsala collection and there is
therefore no reason to assume that this specimen has type status. Specimens which are regarded as having type
status are preserved in the Stockholm collection (Fernholm & Wheeler, 1983) and in the Gronovius collection
(Wheeler, 1958).
Percifoms
Serranidae
Genus and species indet
Pmcu rudulu Syst. Nut. (10): 294 (1758); Syst. Nut. (12): 488 (1766).
Specimen: ZIU 52 [47]; S.L. 149 mm, T.L. 180 mm; in spirit, good condition but faded.
Label: Perca Radula.
Discussion: Percu rudulu was named solely on the earlier description in Arnoenilutes Acudernicue, the reprinted
edition of Balk’s dissertation Museum Adolph-Fridericianum (Linnaeus, 1749a) although the pagination was
cited as 313 in error for 315. The present specimen is labelled Perm Rudulu but the label is not original..
Lonnberg (1896) referred to this specimen saying that it was probably not the type ofP. rudulu as the specimen
does not agree with the description. Despite this there is evidence that a specimen labelled with this name was
in the collection before 1787 (Thunberg, 1787) and in 1828 (Thunberg m catalogue). It is difficult to be
certain if this was the specimen from the Crown Prince’s collection described by Balk. It seems probable that
Lbnnberg was correct in disputing this as recent examination of the dorsal and anal fin elements resulted in
counts o f D XI. 15 and A 111 8, whereas Linnaeus (1758) gave (in modern notation) D XI. 10 and A 111. 10
Linnaeus (1749a) gave D. X. 10 and A 111. 10.
Percidae
Perm Jluviutilis Linnaeus, 1758
Specimen: ZIU 197 1871; S.L. 96 mm, T.L. 11 1 mm; in spirit, good condition.
Label: Perca/fluviatilis/monstrosa/Mus. Linn.
LINNAEAN FISHES IN UPPSALA
171
Discussion: This specimen was listed by Thunberg in his manuscript list of 1828 as specimen “delta, monstrosa
Thunberg”, by Lonnberg (1896: 43) as specimen number 87, and by Holm (1957: 45) as specimen number 197
“Perca Jluuiatilis monstrosa Thunb.”. It is in no way involved in the typification of Linnaeus’s taxon Perca
Jluuiatilis. According to Holm it was part of the Ahlstromer-Linnk donation.
Gymnocephalus cemuus (Linnaeus, 1758)
Specimen: ZIU 200 [91]; S.L. 110 mm, T.L. 131 mm; in spirit, good condition.
Label: Perca/cernua/Mus. Linn.
Discussion: A specimen ofPerca cemua was listed by Thunberg (1787) and in his manuscript catalogue of 1828.
Lonnberg (1896) also listed this as specimen 91 without comment as did Holm (1957) as specimen 200. All
three attributed it to the Ahlstromer-Linnk donation.
There is no evidence that i t is involved in the typification of Perm cemua Linnaeus, 1758.
Echeneidae
Remora remora (Linnaeus, 1758) [Fig. 101
Echeneis remora Syst. Nat. (10): 260 (1758); Syst. Nut. (12): 446 (1766).
Specimen: ZIU 58 [54]; S.L. 126 mm, T.L. 155 mm; in spirit, faded but in good condition.
Label: None.
Discussion: Echenels remora was based on four earlier descriptions, Amoenitates Academicae, the reprinted edition of
Balk’s dissertation Museum Adolpho-Fridericianum (1749a), Artedi ( 1 738), Gronovius ( 1754) and Catesby ( 1743).
The last reference was transferred to Echeneis ncucrates by Linnaeus (1766).
A specimen of Echeneis remora, attributed to the Adolf Fredrik donation was listed by Thunberg ( 1787) and in
his manuscript catalogue of the collection of 1828. This attribution was also given by Holm (1957) as his
number 54. However, Lonnberg (1896) was apparently unable to find this spirit specimen as he referred only
to three dried specimens which he identified as Echeneis neunates (see below), with the result that he inferred
that Linnaeus’s E . remora was synonymous with E. neucrates. The absence of the original label was, no doubt,
the reason for his failing to recognize the specimen as Linnaean. There seems no reason to doubt that it is the
Adolf Fredrik donation specimen and is therefore part of the type series; no other putative type specimen has
survived.
Echeneis neucrates Linnaeus, 1758
E c h e l s neucrates Syst. Nut. (10): 261 (1758); Syst. &at. (12): 446 (1766).
Specimens: ZIU 82; S.L. 190 mm, T.L. 215 mm; whole dry specimen.
Label: Echeneis/Neucrates/Mus Ad Frid.
ZIU 84; S.L. 264 mm; T.L. 300 mm; whole dry specimen mounted upside down.
Figure 10. Remora remora Linnaeus, 1758 (ZIU 58), type specimen.
172
A. WHEELER
Discussion: Echeneis neucrates was based on three sources, a phrase of description and meristic data, and
published references to Vallisneri (1733) and Hasselquist (1757). In Linnaeus (1766) a reference to Catesby
( 1743) was transferred from the account of E. remora in Linnaeus ( 1758). The first of these sources suggests that
the data were derived from a specimen examined by Linnaeus.
Fernholm & Wheeler (1983) concluded that these data may have derived from a Hasselquist specimen
which it had been intended to cite in the second volume of the Mucum Adolphi Fridcrici (Linnaeus, 1764a) but
which was inadvertently omitted. They suggest that a specimen in the Swedish Museum of Natural History
(NRM 3688) may be a Hasselquist specimen and therefore could be a type. This is, however, not certain, and
although the Stockholm specimen is old, dating from the “Garnla samlingen” collection, its early history is not
known.
It seems more probable, now that the Uppsala collection has been properly examined, that the first of the
Linnaean sources may have referred to a specimen in the donation of the Crown Prince Adolf Fredrik (1746) to
the Uppsala Academy collection. If so, then specimen ZIU 82, which is labelled Mus. Ad. Frid., may be the
original specimen examined by Linnaeus.
Although no specimen of EchenCiS ncucrates was described by Balk in his dissertation (Linnaeus, 1746a) on the
donation of Adolf Fredrik, this does not mean that there was no example in the collection. It is accepted that
the museological dissertations described novelties rather than well known species. The first of these specimens
(ZIU 82) is probably a type specimen.
Lonnberg (1896) and Holm (1957) both recorded three dried specimens in the collection.
Carangidae
Trachurus sp. [Fig. 1 I]
Scombcr trachurus Syst. Nat. (10): 298 (1758); Syst. Naf. (12): 294 (1766).
Specimen: ZIU 104 [27]; S.L. 165 mm, T.L. 193 mm; in spirit, but has been dried.
Label: Scomber.
Discussion: Scomber trachurus was based on four earlier published sources Artedi ( 1 738), Linnaeus ( 1 754a),
Hasselquist ( 1757) and Linnaeus ( 1754b). The specimens described by Linnaeus (1 754a) and Hasselquist
( 1 757) may have been identical and are discussed by Fernholm & Wheeler (1983) who concluded that in view
of their curatorial history it was no longer possible to recognize for certain these specimens amongst others in
the Swedish Museum off Natural History collection.
The description by Linnaeus (1754b) was tPe dissertation of Johann Odhelius published under the title
C h i m ‘ a Lugerstromiana which described the donation of Magnus Lagerstrom to the Academy. This was later
reprinted by Linnaeus (1 759) in the series Amomztates Acadmricnc, and this title was cited by Linnaeus ( 1766).
The species numbered 27 in Linnaeus (1 754b) is named Scomber (Trachurus) and has to be considered as part
of the type series. Unfortunately, as the dissertation title suggests, the Lagerstrom donation was of Chinese
origin. The specimen therefore is more probably referable to Tr0churusjaponz’cz.s (Temminck & Schlegel, 1842).
series of
7.frachurus (Linnaeus, 1758);
LINNAEAN FISHES IN UPPSALA
173
However, as the specimen is not in good condition it has not been examined closely so the entry in the present
catalogue appears as Truchurus sp. It is clear though that if this specimen is part of the Lagerstrom donation the
proposal made by Fernholm & Wheeler (1983) to regard it as a putative lectotype is not tenable.
A specimen of Truchurus (as Scomber truchurus) was listed by Thunberg (1787) Lonnberg (1986) and Holm
(1957) as originating in the Lagentrom donation. There seems to be little doubt that these entries refer to the
present specimen.
Selene uomer (Linnaeus, 1758)
Zeus Vomer Syst. Nat. (10): 266 (1758); Syst. Nut. (12): 454 (1766).
Specimen: ZIU 189 [74]; S.L. 109 mm, T.L. 132 mm; in spirit in good condition.
Label: No old label.
Discussion: This specimen was listed by Thunberg (1787), and in his manuscript catalogue of 1828, as deriving
from the Ahlstromer-Linne donation of 1749. Lonnberg (1896) did not list the specimen (according to Holm,
1957, reporting it as number “75 is lost”), but by the time Holm’s catalogue was published the specimen had
been rediscovered.
Zeus uomer was named by Linnaeus (1758) from the description by Patrick Browne ( 1 756) and by Linnaeus
(1754a) in Museum Adolphi Frihrin’. In Linnaeus (1766) the reference to Browne had been omitted, and there
were minor textual alterations and an addition to the description that followed the diagnosis and reference.
The reference to the Museum Adolphi Fridm’ci (Linnaeus, 1754a) therefore assumes increased importance.
Fernholm & Wheeler (1983) reported that there were two specimens of Selene uomer in the Swedish Museum
ofNatural History collection (NRM, LP 17, NRM, LP 17a), both ofwhich were probably part of the material
from the Museum Adolph Fridnici collection, and thus were part of the type series. The present Uppsala
specimen is presumed not to be a type.
Trachznotus ouatus (Linnaeus, 1758) [Fig. 121
Gasterosteus ouutus Syst. Nut. (10): 296 (1758); Syst. Nat. (12): 490 (1766).
Specimen: ZIU 202 [93]; S.L. 129 mm, T.L. 168 mm; in spirit, good condition.
Label: Gasterosteus/ovatus/Mus. Linn.
Discussion: Gasfcrosteus ouatus was described without any reference to earlier literature and must therefore have
been based directly on examination of a specimen. The description was identical in both editions of the Systema
Naturue (Linnaeus, 1758, 1766).
A specimen labelled Gasterosteus ouatus was listed by Thunberg (1787) as being present in the Museum of
Uppsala University having been derived from the Ahlstromer-LinnC donation. The same specimen was
reported by Lonnberg (1896) and Holm (1957). Lonnberg drew attention to the type status of the specimen
and discussed the confused taxonomic history of this species. Wheeler (1963) also discussed the nomenclature of
this fish in relation to the European species of Trachznotus.
There is no doubt that this is the type specimen of Trachinotus OZJQ~U (Linnaeus, 1758).
Figure 12. Truchinotus ouatus (Linnaeus, 1758) (ZIU 202), holotype.
LINNAEAN FISHES IN UPPSALA
175
& Wheeler (1983) found no specimen in the collection of the Swedish Museum of Natural History that could
have represented Hasselquist’s material.
Chaetodontidae
Chuetodon cupistrutus Linnaeus, 1758 [Fig. 131
Chuetodon cupistrutus $st. Nut. (10): 275 (1758); Syst. Nut. (12): 465 (1766).
Specimens: ZIU 51 [GI; S.L. 97, 55 mm, T.L. 116, 66 mm in spirit, condition fair to good.
Labels: Chaetodon capistratus Mus. Ad. Frid.
Discussion: The species Chaetodon cupisfratus was based on three earlier literature sources, Linnaeus ( 1 754a),
Gronovius (1756) and Laurent Balk‘s dissertation Museum Adolpho-Fridericianum which was later reprinted
by Linnaeus (1 749a) in the Amenitufes Acudemicue.
A specimen of this species was listed Thunberg (1787) (and in his manuscript list of 1828), Lonnherg (1896)
and by Holm (1957) as being from the donation of the Crown Prince Adolf Fredrik to the University Museum.
Only Holm commented that there were two specimens (“1 store 0. 1 mindre ex. i alk.”) now in the collection at
Uppsala. Both are labelled as being from the Adolf Fredrik donation, and if these labels are correctly associated
with the specimens then both must be accorded type status.
Fernholm & Wheeler (1983) drew attention to the survival of other putative type specimens in the Swedish
Museum of Natural History (that of Linnaeus (1754a) and in the British Museum (Natural History) (the
Gronovius (1756) specimen).
Chuetodon striutus Linnaeus, 1758 [Fig. 141
Chuetodon striutusSysl. Nut. (10): 275 (1758); Syst. Nut. (12): 464 (1766).
Specimen: ZIU 50 1451; S.L. 124 mm, T.L. 144 mm; mounted half-skin.
Labels: Chaetcdon/striatus/Mus. Ad. Frid. pinned to the specimen.
Discussion: Chuetodon striutus was named by Linnaeus (1758) from four earlier published accounts, namely,
Linnaeus (1754a), Artedi (1738), Linnaeus (1749a) and Gronovius (1754). The specimens described by
Linnaeus (1754a) and Gronovius (1754) still survive in the Swedish Museum of Natural History (Fernholm &
Wheeler, 1983) and in the British Museum (Natural History) (Wheeler, 1958) respectively. The latter is a dry
half-skin.
The present specimen was listed by Thunherg (1787) (and manuscript catalogue of 1828), Lonnherg (1896)
and Holm (1957) as originating in the donation of the Crown Prince Adolf Fredrik in 1746. It was this
collection, or at least the novelties in it, which formed the substance of Laurent Balk’s dissertation, Museum
AdoLpho-Fridericiunum (Linnaeus, 1746a) which was later reprinted in the Amonoenztates Acudemicue (Linnaeus,
1749a). There is therefore every reason to regard it as part of the type series of C. striutur.
There are several other examples where specimens of the same species are present in both the Stockholm and
Uppsala collections, having at some time been part of the Adolf Fredrik collection. The present case is unusual
in that this specimen is a skin, while the Stockholm specimen is in alcohol.
Figure 14. Chuetodon striutus Linnaeus, 1758. (ZIU 50), part of the type series.
176
A. WHEELER
Figure 15. Abudfhf saxatifis (Linnaeus, 1758). (ZIU 49), part of the type series.
Pomacentridae
Abudefduf saxatifis (Linnaeus, 1758) [Fig. 151
Chaetodon saxatifis $st. Nut. (10): 276 (1758); Syt. Nut. (12): 466 (1766).
Specimen: ZIU 49 [44];S.L. 86 mm, T.L. 106+mm; spirit, in poor condition.
Label: Chaetodon/saxatilis/Mus.Ad. Frid.
Discussion: Chuetodon saxatilis was based on three earlier descriptions in published sources, namely Linnaeus
( 1754a), Linnaeus ( 1749a), and Gronovius ( 1754). The Linnaeus ( 1749a) description was that published in
Laurent Balk’s dissertation of 1746, Museum Adolpho-Fridericianum, which concerned the donation of
specimens by the Crown Prince during that year.
Figure 16. Amph$wion polymnw (Linnaeus, 1758). (ZIU 198), holotype.
LINNAEAN FISHES IN UPPSALA
I77
The present specimen was listed by Thunberg (1787), and in his manuscript catalogue of 1828, by Lonnberg
(1896) and Holm (1957) as originating in the donation of Adolf Fredrik. There is thus every reason to suppose
that it is the specimen described by Linnaeus and therefore part of the type series.
The other specimens described by Linnaeus (1754a) and Gronovius (1754) have both survived, one in the
Swedish Natural History Museum, Stockholm, and the other in the British Museum (Natural History) (see
Fernholm & Wheeler, 1983 and Wheeler, 1958).
Amphiprion polymnus (Linnaeus, 1758) [Fig. 161
Percapolymna Syst. N a t . (10): 291 (1758); Syst. Nat. (12): 484 (1766).
Specimen: ZIU 198 [88]; S.L. 102 mm, T.L. 122 mm; in spirit, good condition.
Label: Perca/polymnia/Mus. Linn.
Discussion: Perca polymna was named by Linnaeus (1758) on the basis of a specimen in the collection of Uppsala
University (“Mus. acad.”), no reference to a literary source was given. Later, Linnaeus (1766) added literary
references to his account namely Gronovius (1754) and Seba (1758) but these descriptions are not involved in
the typification of the species.
A specimen of Perca polymnia is listed by Thunberg (1787) as originating in the Ahlstromer-Linnt donation to
the Academy of 1749 and later. The same specimen was listed by Lonnberg (1896) as number 88 of the
Ahlstromer-Linnt donation. As he stated there is no reason to doubt that this was the type-specimen of Perca
polymna. The specimen was again fisted by Holm (1957).
.
This is the type specimen ofAmphiprionpolymnus (Linnaeus, 1758). Despite doubt expressed by Lonnberg that
this fish was referable to A . QifasciatusBloch & Schneider, 1801, it is certainly referable to A . polymnus Linnaeus,
1758) as defined by Allen ( 1972).
Cepolidae
Cepola rubescens Linnaeus, 1766
Cepola taenia $st. Nat. ( 12j : 445 ( 1766).
Cepola rubescens Syst. N a t . ( 12): 445 ( 1766).
Specimen: ZIU 182 [67]; S.L. 183 mm, T.L. 214 mm; in spirit, moderate condition.
Label: Cepola/taenia/Mus. Linn.
Discussion: The taxonomic history of this species was discussed by Fernholm & Wheeler (1983) and reference
was made to this specimen in that paper. To summarize, the name C. rubescens was based solely on the
description in Linnaeus (1764a) of one or more specimens in the collection of King Adolf Fredrik; a specimen
which may have been in that collection is still preserved in the Swedish Museum of Natural History,
Stockholm. Simultaneously with the publication of that name, Cepola taenia Linnaeus, 1766 was proposed. The
only literary reference cited was to Artedi ( 1 738), but unusually for species of fish named in the Systema Naturae
there is a paragraph of description which suggests (by the inclusion of details not in Artedi) that Linnaeus had
examined a specimen which he identified with Artedi’s Taenia.
The collection of fishes in Uppsala contained specimens identified as both Cepola tamin and C. rubescens in the
donation of Ahlstromer-Linnt (Thunberg, 1787). The enumeration of the collection in systematic order
(compiled by Thunberg in 1828) complicates this simple situation because it lists specimens of C. rubescens from
both the Ahlstromer-Linnt and the Thunberg collections, as well as a specimen of C. taenia from the Adolf
Fredrik donation. However, Lonnberg (1896) listed only a single specimen of Cepola taenia from the
Ahlstromer-Linnt donation, while stating that the specimen of C. rubescenr was lost. (He made no mention of
the Thunberg specimen which lay outside the terms of reference of his paper on Linnaean types.)
The situation was made more complicated still by Holm (1957) who wrote of Lonnberg’s number 66 Cepola
taenia that it was missing (contradicting the latter’s statement that it was present) and listed this specimen
(Lonnnberg’s number 67) as present with a Thunberg label C. taenia.
This does not seem to shed much light on the question as to whether this specimen is the type of Cepola taenia.
The early label with it is in Thunberg’s hand and is therefore post-Linnaean. Thunberg’s (1828) catalogue of
the collection lists only one C. taenia and that from the donation of Adolf Fredrik. If this catalogue is accurate
then the present specimen cannot be the type specimen. However, if it is erroneous on the origin of the
specimen then it may be a type specimen of C. taenia. It is certainly not the type of C. rubescens.
Lutjanidae
Luijanus sp.
Pentapus paradiseus Gunther, 1859.
Specimen: ZIU 223; S.L. 115 mm, T.L. 187 mm; spirit in good condition.
Label: No original label (recent internal label Pentapus paradiseus Gunth. Mus. Linn.).
Discussion: This specimen was listed by Holm (1957) as Polynemzs paradiseus; it is not a member of the family
Polynemidae. Its identification is uncertain but at some time it has been identified with Gunther’s later
described species Pentapus paradiseus. I am not aware of any Linnaean name which could relate to this species
and it seems to have no type status.
A. WHEELER
178
Linnaeus, 1758.
Labridae
Ccntrolabm exolefur (Linnaeus, 1758) [Fig. 171
Specimen: ZIU 194 [82]; S.L. 99 mm, T.L. 117 mm; spirit, in moderate condition.
Labels: Original label Labrrcs/chinensis/Mus. Linn. (recent internal label Ctenolabrus exoletus L. = Labrus
chinensis. Mus. Linn.).
Discussion: This specimen bears a Thunberg label attributing to it the name Lubrus chinensis Linnaeus, 1758.
This taxon was named by Linnaeus evidently from a specimen (in the absence of any literary reference or
citation of a specimen in another collection) with the locality “Habitat in Asia”. The diagnosis of the species is
brief “Labrus pinna dorsi ramentacea, corpore livido, vertice retuso” but with the anal fin ray count of five
spines in a total of twelve elements suggests that it may have referred to this specimen.
A specimen labelled Lobrus chinnrris is recorded in the Uppsala collection in the list compiled in 1780-81
(Holm, 1957), in Thunberg’s published account of 1787, and in his manuscript list of 1820. Lonnberg (1896)
examined the specimen and identified it as L a b w exolefus ( = Cmtrdubrus exoletus) and suggested that the
Linnaean name L o b m chinnrsis was a junior synonym of L. exoletus. The possibility that at some time in the past
a specimen of C. exolefus had been substituted for the original specimen of L a b m c h i n m i remains, but there is
no evidence to support it, and as Linnaeus’s name L. c h i k does not appear to have been used by later
workers is no purpose in pursuing the suggestion.
I t is, however, curious that Linnaeus should have named this wrasse both as Lubrus exoletus and L. chinensis in
adjacent entries in the &stemu Nafurac,and paradoxical that while the original specimen of L. chine& appears
to have survived that of L. exoletus seems to have disappeared. If it became necessary in a revisionary study to
designate a neotype for L. exoletus it might be desirable to designate this specimen, which is apparently the type
of L. c h i n m s , which name could then bcome a junior objective synonym.
Crenilabrus melops (Linnaeus, 1758)
Specimen: ZIU 192 [80]; S.L. 129 mm, T.L. 161 mm; spirit, in moderate condition.
Labels: Original labels Labrus/melops/Mus. Linn.
Discussion: Linnaeus (1758) named L a b m melops from a draft description which was only published some years
later (Linnaeus, 1764a). The specimen described, which Fernholm & Wheeler (1983) considered to be the type
of the species, is preserved in the Swedish Museum of Natural History, Stockholm. There is therefore no reason
to assume that the Uppsala specimen is a type.
Nevertheless it appears to be a specimen from Linnaeus’s collection as there is an entry under the name
Lubrus melops in the manuscript catalogues of the collection made in 1780-81, and 1820, as well as in the lists
published by Thunberg (1787) and Lonnberg (1896). AU of these except for the first give the origin of the
specimen as the donation of Ahlstrijmer & Linne.
LINNAEAN FISHES I N UPPSALA
179
Figure 18. Labrus mixtus Linnaeus, 1758. ( Z I U 193), holotype of Labrus ossifragus Linnaeus, 1758
Labrus mzxtus Linnarus, 1758 [Fig. 181
Labrus ossifagus Syst. N a t . (10): 286 (1758); Syst. N a t . (12): 478 (1766).
Specimen: Z I U 193 [81]; S.L. 264 mm, T.L. 309 mm; in spirit good condition.
Labels: Labrus/ossifragus/Mus. Linn.
Discussion: The name Labrus ossifagus was published by Linnaeus (1758) based apparently on a specimen
available to him. The present specimen appears to have a continuous curatorial history in the Uppsala
collection as it was listed in the manuscript catalogue compiled in 1780-86 only two years after Linnaeus’s
death, in Thunberg’s ( 1 787) catalogue, as well as in his manuscript catalogue of 1820, and Lonnberg’s (1896)
study of the collection. T h e three last entries all refer the specimen to the Ahlstromer-Linni: donation of 1749.
There seems to be little doubt that this is the type specimen of Labrur oss2fagu.s Linnaeus, 1758.
Lonnberg (1896) pointed out that Thunberg in his listings of the collection had corrected Linnaeus’s name
ossifgus to ossifragus, the latter being undoubtedly the intended spelling.
Parenthetically it may be noted that Linnaeus named this species three times in the tenth edition of the
Systema Naturae (as Labrus bimaculatus, L . mixtus, and L . ossifragus).
Although some authors have recently used the name Labrus bimaculatur (see Bauchot & Quignard, 1973),
others have chosen to employ L . mzxtur on the grounds of its much greater usage.
Hemipteronotus sp. [Fig. 191
Coryphaena uirens $st. N a t . (12): 448 ( 1766).
Specimen: Z I U 183 [68]; S.L. 169 mm, T.L. 190 mm; in spirit in good condition.
Label: Coryphaena/virens/Mus. Linn.; recent label Novacula cultrata C & V = Coryphaena virens Linn.
Discussion: Coryphaena uzrens was described by Linnaeus (1766) with the locality Habitat in Oceano Asiatico. A
specimen with this name was listed in the collection in Uppsala in the catalogue published by Thunberg
( l i 8 7 ) , in the manuscript list of 1820, by Lonnberg (1896) and by Holm (1957). A specimen of Coryphaena
pompilus? is listed in the 1780-81 catalogue and this note of interrogation may refer to doubt as to it being
conspecific with C. pompilus Linnaeus. This seems possible as the list of 1820 refers the only specimen of
C. pompilus to Thunberg’s donation which was made in 1775 and the published catalogue of 1787 lists simply
C. uirens. As there appears to be a continuous curatorial record for this specimen I have little doubt that it is the
type of Coryphaena uirens.
Lonnberg (1896) appears to have reached the same conclusion but was somewhat less positive; “it is
uncertain, although probable, if it is a type-specimen”.
T h e name Coryphaena uirms was used by Gmelin ( 1 789) and was also employed by Solander for a specimen of
Hemipteronotus ( = Nouacula) on Cook‘s first voyage (Wheeler, 1986). From the drawing (and possibly the
manuscript) associated with this voyage Valenciennes (1839) published Xyrichthys uirens, attributing the trivial
epithet to Parkinson. Neither Valenciennes (1839) nor Wheeler (1986) realized that it was a Linnaean taxon.
180
A. WHEELER
If Linnaeus’s note on the geographical origin of this specimen (“Ocean0 Asiatico”) is correct then it is
unlikely to be Nouacula cultrata Cuvier & Valenciennes, 1839 (a junior synonym of Hemipteronotus nouacula
Linnaeus, 1758 (Randall, 1965)). For this reason I have not attempted to identify the specimen to species, nor
do I favour the adoption of Liinnberg’s (1896) suggestion that the Atlantic-Mediterranean species should be
called Nouaculu M’rm. I t is possible that this Linnaean name should be regarded as a nomen oblitum, as it appears
not to have been used since the eighteenth century.
7hularsoma lunare (Linnaeus, 1758)
Labrus lunaris Syst. Nat. (10): 283 (1758); Syst. Nat. (12): 474 (1766).
Specimen: ZIU 195 [83]; S.L. 139 mm, T.L. 172 mm; in spirit, in good condition.
Label: Sparus honoratus Mus. Linn. Sparus honoratus Mus Thun.; recent label Julis lunaris.
Discussion: As Lonnberg (1896) pointed out there has been a confusion with labels between this specimen and
that of Sparus honoratus. Spurus honoratus was named by Thunberg (1787) with a short account of the meristic
characters of the specimen in the Ahlstromer-LinnC donation. According to this listing there should also have
been a specimen of Labrus lunaris in the same donation.
Labrus lunaris was named by Linnaeus ( I 758) from a specimen he had examined and a specimen described by
Gronovius ( 1 756). The latter specimen is a dry skin now in the British Museum (Natural History). An entry for
both Labrus lunuris and Sparus honoratus appears in Thunberg’s (1 787) list of the collection and also in the 1820
manuscript list. By the time Liinnberg (1896) published his list there was only a single specimen of Labrus lunaris
labelled as Sparus hmorutus, suggesting that the change of labels had occurred during this interval. However, it
seems very probable that this specimen is the other type of Thdassoma lunare (Linnaeus, 1758), even though the
change of labels casts a slight element of doubt on its status.
Ctenolabrus rupestris (Linnaeus, 1758)
Specimen: ZIU 196 [85]; S.L. 126 mm, T.L. 149 mm; in spirit.
Labels: Sciaena/umbra/Mus. Linn.
Discussion: This specimen was listed by Holm (1957) as originating in the Ahlstromer-LinnC donation.
Lonnberg (1896) similarly attributes it to this donation but added, “By some mistake Thunberg has labelled a
specimen of Ctenolabrus rupestris (Linnaeus) with the above [Sciaena umbra] name”.
There is no question of this specimen being involved in the typification of SCiaeM umbra which was named
from the description by Artedi (1738) and from the account in Hasselquist (1757). No material from Artedi‘s
descriptions still exists, and such specimens from Hasselquist’s collections as exist are in the Swedish Museum of
Natural History, Stockholm (Fernholm & Wheeler, 1983); these authors do not list the specimen of Sciamn
umbra described by Linnaeus. There is no question of this being a type specimen.
LINNAEAN FISHES IN UPPSALA
181
Trachinidae
Trachinus draco Linnaeus, I758
Specimen: ZIU 174 1551; S.L. 227 mm, T.L. 277 mm; dry, mounted on a board.
Label: Trachinus/Draco./Mus. Linn.
Discussion: This specimen was listed as being part of the Ahlstromer-Linni: donation of 1749 and later years by
Thunberg (1787), Liinnberg (1896) and Holm (1957). At no time has it been recognized as type material.
Truchinus draco was named from earlier accounts by L. Gronovius (1754), J. F. Gronovius (1742), Artedi
( I 738) and Linnaeus ( I 751). There is no evidence that this specimen can be associated with Linnaeus's (1 751)
description.
Uranoscopidae
L'ranascopus scaber Linnaeus, 1758
Specimen: ZIU 190 [76]; S.L. 162 mm, T.L. 206 mm; in spirit; good condition, thread at mouth to allow
suspension of fish.
Label: Partly destroyed.. .opus/. . .ar &/Mus. Linn.
Discussion: This specimen was listed by Holm (1957) (and earlier authors) as being part of the
Ahlstromer-Linnk donation of 1749. At no time has it been claimed to have been part of the type series.
Discussion of the typification of this taxon was given by Fernholm & Wheeler (1983); they concluded that
the only extant type specimen was in the Swedish Museum of Natural History, Stockholm.
Zoarcidae
Zoarces viuiparus (Linnaeus, 1758)
Specimen: ZIU 180 1631; S.L. 268 mm, T.L. 270 mm; in spirit; good condition.
Label: Blennius/viviparus & / M u Linn.
Discussion: This specimen was listed by Holm (1957) and by earlier authors, as belonging to the
Ahlstromer-Linnt donation of 1749 and later. At no time has it been considered to be a type specimen. The
taxon Blennius vzviparus was named from five earlier accounts, viz. Artedi (1738), Linnaeus (1754a), L. T.
Gronovius (1754), J. F. Gronovius (1742) and Gissler (1748) (as Act. Stockh. 1748). None of the specimens
described by these authors is likely to be in Uppsala; that described by Gronovius (1754) (which is possibly the
same as that described by J. F. Gronovius) is believed to be in the British Museum (Natural History) (Wheeler,
1958).
Clinidae
Clinus superciliosus (Linnaeus, 1758) [Fig. 201
Blennius superciliosus Syst. Nat. (10): 257 (1758); Sysf. f l a f . (12): 442 (1766).
Specimen: ZIU 54 [50]; S.L. 107 mm; T.L. 115+damage mm; in spirit; moderate condition.
Label: No original label.
Discussion: The name Blennius supercilzosus was based on two earlier descriptions. The first cited (as Amoen. scad.)
was the dissertation defended by Laurentius Balk (Linnaeus, 1749a) in which specimens from the donation of
the Crown Prince Adolf Fredrik to the University in 1745 were described. The second was Gronovius's (1 756)
description in the Museum Jchthyologicum. Wheeler (1958) drew attention to three specimens of this species in the
Gronovius collection but was doubtful whether they were the originals of his 1756 description and thus part of
the type series. This should, however, be re-examined.
As the specimen described by Linnaeus (1 749a) was from the donation of the Crown Prince to the University
of Cppsala it would he expected to be found in the present collection. Even though it has no label there is every
reason for thinking that it is part of the type series of Blennius superciliosus. It was recognized as a type by Holm
(1957) hut not by Liinnberg (1896). However, a specimen of Blennius superciliosus has been listed in successive
catalogues of the collection from 1787, and there is little doubt that this is the specimen described in Balk's
dissertation.
Blenniidae
f'arah/ennius cornulus (Linnaeus, 1758) [Fig. 211
Blennzus cornutus *sf. N a t . (10): 256 (1758); Syst. N Q ~(12):
.
441 (1766).
Specimen: ZIU 53 [49]; S.L. 89 mm, T.L. 105 mm; in spirit; moderate condition.
Label: Blennius/cornutus./Mus. Ad. Frid.
Discussion: The name Blennzus carnutus was based on two earlier descriptions of which only one was published
before 1758. They were Amoen. acad. ( I , p. 316) and Mus. A d. Fr. 2, which was not published until 1764. The
Amoenitates Academzcae refers to the dissertation of Laurent Balk (Linnaeus, 1749a) in which specimens from the
182
A. WHEELER
donation of the Crown Prince Adolf Fredrik were described. The second reference was to the then unpublished
Museum Adolghi Friderin’ Volume 2, describing specimens in the King’s collection (Linnaeus, 1764a). There is no
doubt that the two specimens were originally a single lot in the royal collection.
A specimen recognized as Blcnnius cornutuc was listed in the Academy collection, by Thunberg ( 1787) and in
his unpublished list of 1828, by Lonnberg (1896) and by Holm (1957) who recognized it as a type specimen. As
no material of this species was found which could be attributed to the King’s collection in Stockholm
(Fernholm & Wheeler, 1983) this present specimen appears to be the only surviving type material of Blennius
comutus.
Pholidae
Pholis gunnellus (Linnaeus, 1758)
Blmniusgunncllus *st. Nut. (10): 257 (1758); Syst. Nut. (12): 443 (1766).
Specimen: ZIU 179 [62]; S.L. 147 mm, T.L. 157 mm; in spirit.
Label: No original label.
Discussion: This specimen was listed by Holm (1957) as being present in the collection and part of the
Ahlstromer-Linnt donation of 1749. A specimen of this taxon was reported by Thunberg (1 787) and in an
unpublished list of 1828, but Liinnberg claimed that the specimen (his number 62) was lost.
As the species was named on the basis of descriptions in the Museum Ado&hi Frideriium (Linnaeus, 1754a) and
Artedi (1738) there is no reason to expect his to be a type specimen. The only existing type material of this
taxon is in the Swedish Museum of Natural History, Stockholm, (Fernholm & Wheeler, 1983). This was the
specimen described by Linnaeus (1754a).
Ammodytidae
Ammodytur tabianus Linnaeus, 1758
A m d y # c s tobinnusSyst. sat. (10): 247 (1758); Syst. Nut. (12): 430 (1766).
Specimens: ZIU 173 [54]; S.L. 151, c. 152 mm, T.L. 160, 161 mm; dried, the larger one broken in middle.
Label: Ammodytes/tobianus/Mus. Linn.
Discussion: The name Anunodytcs tobianuc was based by Linnaeus on five earlier descriptions, Artedi (1 738) and
Linnaeus (174613, 1751, 1745, and 1754a) in that order.
A specimen named as this species, preserved in the Swedish Museum of Natural History, Stockholm
(Fernholm & Wheeler, 1983), is referable to A . mrinus Raitt, 1934. A second specimen is preserved in the
Linnean Society of London (Wheeler, 1985), which is thought to be part of the type series having possibly been
collected during Linnaeus’sjourney to Oland and Gathland.
LINNAEAN FISHES IN C‘PPSALA
I83
Figure 21. Parablennius comutus (Linnaeus, 1758), (ZIU 53), part of the type series
The Uppsala specimens, having originated as Linnaeus’s donation, may have been collected by Linnaeus on
this journey (Linnaeus, 1745), but there is no proof of this. If it were provable then they would be part of the
type series.
Gobiidae
Gubius niger Linnaeus, 1758
Gubius niger $st. Nat. (10): 262 (1758); Syst. Nat. (12): 449 (1766).
Specimen: ZIU 184 [69]; S.L. 122 mm, T.L. 148 mm; in spirit good condition.
Label: Gobius/niger a/Mus. Linn.
Discussion: The name Gobiur niger was based by Linnaeus on three earlier literary references and to a single
unattributed diagnosis. In Linnaeus (1766) this last is referred to the second volume of the Museum Adolphi
Friderici (Linnaeus, 1764a). In view of these references there is little possibility of the Uppsala specimen being
part of the type series.
This specimen is, however, associated with the Ahlstromer-Linnt donation of 1749 and is listed as such
Holm (1957), Lonnberg (1896), Thunberg’s manuscript catalogue of 1828, and Thunberg (1787).
Boleophthalmus pectinirostris (Linnaeus, 1758) [Fig. 221
Gobiuspectinirostris Syst. Nm. (10): 264 (1758); Syst. Nat. (12): 450 (1766).
Specimen: ZIU 106 [29]; S.L. 83 mm, T.L. 100 mm; in spirit, condition fair.
Label: Gobius/pectinirostris/Mus. Lagerstr.
Discussion: The taxon Gobius pecfznirostrir was based by Linnaeus (1 758) on Odhelius’s dissertation Chinensia
Lagerstromiana (Linnaeus, 1754b) and Osbeck’s ( 1 757) description. The present specimen is clearly that referred
to by Linnaeus (1754b) as the diagnosis “dentibus maxillae infenoris horizontalibus” describes the dentition of
this specimen.
‘This is another example of an Uppsala specimen being described in the Odhelius dissertation and by Osbeck.
This strengthens further the case made that the Lagerstrom donation contained specimens collected by Osbeck.
A specimen of Gobius pectinirostris was listed by Thunberg ( 1 787), and in his manuscript catalogue of 1828,
both cited as from the Lagerstrom donation. Lonnberg (1896) also listed a specimen under this name but
because Linnaeus had cited a reference to Osbeck‘s “Apocryptes chinensis” he concluded that the correct name
to employ was Boleophthalmus chinensis (Osbeck & Linnaeus). This view is not sustainable in modern taxonomic
practice.
Holm (1957) listed this specimen, adding in his remarks-one large example in alcohol Type? (Lonnberg),
but he also referred to specimen 105 using the same name Gobiuspectinirostris, and adding-one small example
in alcohol Type? (Lonnberg). This confusion is apparently of long standing for Thunberg (1787) listed Gobius
I a4
A. WHEELER
LINNAEAN FISHES IN UPPSALA
185
186
A. WHEELER
pectinirostris with a footnote saying “Gobius eleotris ad duplicates numerandus”. This specimen, however, is not
referable to Boleophthalmus.
I am grateful to Dr P. J. Miller for confirming the identity of this specimen with Boleophthnlmus pectinirostris
(Linnaeus, 1758) of Koumans (1953).
Rhtnogobzus sp. [Fig. 231
Gobius eleolns Syst. #at. (10): 263 (1758); Syst. Nut. (12): 449 (1766).
Specimen: ZIU 105 [29]; S.L. 58 mm, T.L. 70 mm;in spirit, fair condition.
Label: Gobius/pectinirmtris/Mus. Lagerst. (See discussion.)
Discussion: Gobius eleotrir was bornl mi four earlier descriptions, th
Odhelius’s dissertation Chinensin
to the reprinted edition of Balk’s
Lagerstrontanu (Linnaeus, 1754b), the second to Osbeck (1757),
dissertation Museum A
a b l ~ (Linnaeus,
F ~ 1749a), and
T. Gronovius (1756). This was
an unusual combination of sources except that once again citation of Chinarcia Lagm&miana coincides with
Osbeck (1757), this supporting the suggestion that some of Osbeck’s material fomed part of the Lagerstrom
donation. However, the specimen described in the Balk dissertation (which was originally published in 1746)
cannot have originated from osbeck; it was presented to the Academy in 1745 with other specimens, by Adolf
Fredrik, then Crown Prince. No specimens of this species could be located in the Gronovius collection by
Wheeler (1958, 1989).
One immediate problem in recognizing this specimen as the type of Gob& eleohis is the label “Gobzus
pectznzrostrir” in Thunberg‘s hand. In Thunberg (1787) this latter taxon is listed with a footnote “Gobius eleotris
ad duplicates numerandus” (the original specimen of G.ptcriRiresais [ZIU 1061 is listed under the donation of
Magnus Lagerstrom). However, iahis manuscript list of the Uppsala d e c t i o n prepared in 1828, Thunberg
lists only a single specimen of Gebius~tisirohisattributed to bgcmtrijm, but three specimens of G. elotris (sic),
one originating in the donationsdAdodfFndrik (a) and two fmm Thunbcrg (B and 7). The first of these must
be presumed to have been a
n of G.clcohis. Liinoberg (1896)dairncd that specimen number 42 of
was lost, and this is repeated by Holm (1957). It has therefore to be
Balk‘s dissertation (Li
ISfmm the Lagemriim donation. Unnberg (1896) refers to a specimen of
assumed that the presen
Gobius kept “with another fish in a receptacle with Thunberg‘s lab& “Mus. Lageystr”. This suggests that the
two gobiids (105 and 106) were the same bode in 1896 and chat subsequently they have been separated and
the labels (one is much older and kcas legible than the other) have been reallocated to the two specimens. (Dr
Lars Wallin inform me thet tke two b o t h are semi-modern, i.e. do not date &om Thunberg’s time.)
It has to be presumed thcrcfore that this specimen is the missing Lageratrim donation specimen referred to
by Linnaeus (1758) and that while the species name on it is incorrect the attribution of its source to this
donation is correct. If this is so it is part of the type series of Gobiw dcotris Linnaeus, 1758.
I am indebted to Dr P. J. M i e r who examined the specimen and identifies it with the genus Rhznogobius.
Tunioides anguillaris (Linnaeus, 1758) [Fig. 241
Gobius anguillaris Syst. &at. (10): 268 (1758); Sysf. Nut. (12): 450 (1766).
Specimen: ZIU 185 [71]; S.L.240 mm, T.L. 251 mm; in spirit in poor condition.
Label: Gobius anguillaris/Mm. Linn.
Discussion: Gobius anguillmis was described by Linnaeus (1758) from a specimen in his own collection (as shown
by there being no references to literary sources). The locality given for the species is China. There is no doubt
that this specimen is the type material of that taxon as there is a complete and unequivocal curatorial history
for it.
A specimen of Gobius Mguillaris is listed by Thunberg (1787) as present in the Academy Museum from the
Ahlstromer-LinnC donation of 1749 and subsequent years; it is also listed in Thunberg’s manuscript list of
1828. Lonnberg (1896) also listed it and drew attention to the fact that it is referable to Amblyopus caeculus
Cuvier & Valenciennes (correctly A. c d u s (Bloch & Schneider, 1801), pointing out that the Linnaean name
had priority. Holm (1957) also listed it as prcsent in the collection.
The Linnaean name was employed as the senior synonym by Koumans (1953) in the combination Tueniozdes
anguillans.
Trichiuridae
Eupleurogrammus muticus (Gray, 1831)
Specimen: ZIU 172 [53]; T.L. 491 mm; in spirit, fair condition.
Label: No old labels.
Discussion: The possibility that this specimen is part of the type series of Trichurus le#turur Linnaeus, 1758 was
discussed by Lonnberg (1896). He concluded that the name might have been composite, evidently regarding
this as a specimen involved in the typification of the taxon. He did not however claim this specimen to be a
Linnaean type, as Tucker (1956) alleged. Fernholm & Wheeler (1983) in discussing the typification of
T.lcpturus, concluded that there is only one surviving putative type specimen, that which was described by
Linnaeus (1754a) in the Museum Adorphi Fridcrin’ and is now in the Swedish Museum of Natural History,
Stockholm.
187
Figure 24. Tuenioides unguilluris (Linnaeus, 1758), (ZIU 185) holotype.
The present specimen was part of the Ahlstromer-Linnk donation of 1749 and later years (Thunberg, 1787;
Lonnberg, 1896; Holm, 1957) and is undoubtedly of Linnaean origin, but there is no evidence that it was
referred to by Linnaeus in 1758.
The present identification should be regarded as tentative.
Scombridae
Scomber scombrus Linnaeus, 1758
Scomberscombw Syst. Nut. (10): 297 (1758); Syst. Nut. (12): 492 (1766).
Specimen: ZIU 203 [94];S.L. 244 mm; T.L. 275 mm; in spirit, fair condition but bleached.
Label: “Scomber scomber & Mus. Linn” (internal label).
Discussion: The name Scombcr scombrus was based on earlier references to Linnaeus (1 746b), Artedi (1 738) and
Gronovius (1754). None of these would have been based on the specimen in Uppsala.
This specimen is part of the Ahlstromer-Linnt donation of 1749 and later, and is recorded as such by
Thunberg (1787), Ldnnberg (1896) and Holm (1957). I t is not a type specimen.
Monodactylidae
Monodactylus urgmteus (Linnaeus, 1758) [Fig. 251
Chaetodon urgmteus Syst. Nut. (10): 272 (1758); Syst. Nut. (12): 461 (1766).
Specimen: ZIU 103 [26]; S.L. 86 mm, T.L. 110 mm; in spirit, fair condition.
Label: Chaetodon/argentatus/Mus. Lagerstr.
Discussion: The name Chattodon urgmtcus was based by Linnaeus (1758) on the description in Odhelius’s
dissertation Chincnsiu Logcrstomiunu (Linnaeus, 1754b), although later Linnaeus (1766) cited the reprint of this
work in Amomitutes Acudcmicoc (Linnaeus, 1759). The present specimen has a clear curatorial history in the
collection at Uppsala having been cited as from the Lagerstrom donation of 1748 by Thunberg (1787; and
1828 manuscript list), by Lonnberg (1896) and by Holm (1957). The last author questioned its type status but
there can be no doubt that it is the type specimen of C. urgentatus and has the standing of a holotype.
Channidae
Chunna asiatica (Linnaeus, 1758) [Fig. 261
Gymnotusasiaticur $st. Nut. (10): 246 (1758); $56. Nut. (12): 428 (1766).
Specimen: ZIU 171 [52]; S.L. 175 mm, T.L. 210 mm; in spirit, fair condition.
Label: No old label.
Discussion: Gymnotus asiuticur was described in detail in Linnaeus (1758), this description being amplified
slightly in Linnaeus (1766). In neither work did he cite any published sources and it is clear that he was basing
his description on a specimen. A specimen bearing this name was listed from the Ahlstromer-Linnk donations
188
A. WHEELER
Figure 25. Maodocrylur urgmkus (Linnaeus, 1758) (ZIU 103), holotype.
Figure 26. Chunnu asiariCa (Linnaeus, 1758), (ZIU 171), holotype.
of 1749 and later, by Thunberg (1787) (and in his manuscript catalogue of 1828) although there it was listed as
Curapo asialicur. Liinnberg (1896) also listed it and discussed its identification as Chunm;he drew attention to the
note added by Linnaeus (1 766) “dubii Generis, an Anarchichai” in which he indicated his uncertainty about
the generic status of this species. Liinnberg did not doubt that this specimen was the type of Gymnotus &ticus
( = Chunm .riotica) but Holm (1957) failed to acknowledge its type status.
There is, however, no doubt that this is the holotype of Chunnu d c u (Linnaeus, 1758).
LINNAEAN FISHES IN UPPSALA
I89
Callionymidae
Cullionpus bra Linnaeus, 1758
There are two specimens of this species (S.L. 40,39 mm, T.L. 50,48 mm) contained within the bottle which
holds the specimen of MyoxoccpMus quadricomis (ZIU 187). They were mentioned, but not identified, in Holm’s
(1957) list.
Pleurmctifoormcs
Pleuronectidae
Platichthysflcsus (Linnaeus, 1758)
Pleurrmectcsfksus Syst. Nat. (10):270 (1758); syst. NQf.(12): 457 (1766).
Specimen: ZIU 191 [77]; S.L. 140 mm, T.L. 171 mm; in spirit, good condition.
Label: Pleuronectes/Flesus a/Mus. Linn.
Discussion: Pleuroncctcsflcsus was based on five earlier accounts or descriptions, two of which were to different
parts of Artedi (1738), the third to Gronovius (1754) and the remainder to Linnaeus (1746b, 1751). Specimens
which are probably or certainly the originals of descriptions of species in both these works, e.g. Raniceps runinus
and Pelecus cultratus, are preserved in this collection. I t is therefore possible that this specimen is the original of
those referred to in either of Linnaeus’s works cited above. However, with such a common fish in Swedish
waters, this specimen may have been obtained at any time.
This specimen was listed as originating in the Ahlstromer-Linnk donation by Thunberg (1787) and in his
1828 manuscript catalogue, by Lonnberg (1896) and Holm (1957). None of them made any comment
concerning its possible type status, which suggests in the case of Lonnberg, at least, that he felt that it was
unlikely.
Tetraodontijomes
Balistidae
Balistes sp.
Bulistes monoceros +st. Not. (10): 327 (1758); +st. Nut. (12): 404 (1766).
Specimen: ZIU 164 [41]; S.L. 68 mm, T.L. 68 mm+damage to tail; whole dried specimen mounted on block.
Labels: Balistes Monoceros/Mus. Linn. and Balistes ringens Mus. Linn.
Discussion: The old label on this specimen identifies it with Bdistes monoceros Linnaeus, 1758 (now Aleuteres
monoceros) but it was later re-labelled as Balistes ringenr (in 1808 according to Holm, 1957). None of the authors
writing about the Uppsala collection seem to have considered it as possible type material of B . monoceros, a
species which was based on earlier descriptions by Linnaeus (1764a), Osbeck (1757) and Catesby (1743). I
have no supplementary information leading me to suggest that it might have been part of the type material. It
originated in the Ahlstromer-Linnt donation of 1749 and later.
Bulistes vetula Linnaeus, 1758 [Fig. 271
Bulistes vetula S’st. Nut. (10): 329 (1758); +sl. Nut. (12): 406 (1766).
Specimen: ZIU 101 [223; S.L. 195 mm, T.L. 285 mm; dried, stuffed and mounted.
Labels: Balistes vetula/Mus. Lagerst. (two labels).
Discussion: Bulistes vetula was founded on a specimen available to Linnaeus (as shown by a series of meristic
data without reference to a literary source), and to descriptions by Catesby (1731-43), Artedi (1738), Osbeck
(1 757) and Browne (1 756).
The first of these (the meristic data) may have been derived from the specimen in the Academy collection
which was part of the Lagerstrom donation of 1748, as the meristic characters are with one exception identical
with Odhelius’s description in Linnaeus (1754b). This species is there named Balistes capistratos although this
was changed in Linnaeus (1759) to B. uetulu.
This is another example of a specimen collected by Osbeck which is believed to be identical with the
specimen donated to the Academy by Lagerstrom and described by Odhelius (Linnaeus, 1754b).
This specimen was part of the Lagerstrom donation of 1748 and is listed as such by Thunberg (1787) who
noted that it was “siccata”. However, it was not listed in his manuscript catalogue of 1828 and Lonnberg
(1896) reported the specimen of B. vetula as “lost”. However, by the time Holm (1957) catalogued the
collection it had been rediscovered and he listed it and published a photograph of this specimen. Holm
suggested that it was a type, and there seems little doubt that it is the specimen described by Linnaeus
(1754b)-although this reference was not cited as a source in Linnaeus (1758). This specimen should therefore
be accorded type status.
Ostraciontidae
LuctophTs triqueter (Linnaeus, 1758)
Ostrucion triqueter Syst. Nut. (10): 330 (1758); Sysf. Nat. (12): 407 (1766).
Specimen: ZIU 165 (labelled 163 in error) [42]; S.L. 90 mm, T.L. 110 mm; dried and mounted skin.
190
A. WHEELER
Label: Ostracion/triqueter/Mus.Linn.
Discussion: This specimen was part of the donation of Ahlstromer-LinnC of 1749 and subsequent years. It is
listed in the catalogues of the collection issued by Thunberg (1787) and the unpublished list of 1828, and by
Holm (1957), but Liinnberg (1896) noted that it was lost (his number 42). Each of these authors related it to
this donation.
However, the Linnaean name Ostracion triqueter was based on a reference to Linnaeus (1754a) and to Artedi
(1738). The specimens from the former source are, where they exist, in the Swedish Museum of Natural
History, Stockholm (no Ostrncion specimens were listed by Fernholm & Wheeler, 1983), and no specimens
examined by Artedi seem to have survived. There seems therefore no possibility that this is part of the type
series.
“Ostracion” sp.
Ostracion cornutus Syst. Nut. (10): 331 (1758); Syst. Nut. (12): 409 (1766).
Specimen: ZIU 58 ’‘Museum Gustavi Adolphi” donation; S.L. 130 mm, T.L. 165 mm; stuffed, whole fish
mounted.
Labels: Ostracion cornutus/Mus. Gust. Ado. [b has been added later following the word Mus.], another label
on the mounting board reads Ostraciotl diaphanus B1. Schn.
Discussion: According to Holm (1957) this specimen originated in the donation of King Gustav Adolf
(Gustav IV) who presented to the University the remains of the collection which had been made by his
grandmother Queen Louisa Ulrika, at the castle of Drottningholm. Most of the fishes in this collection
preserved in alcohol were removed to the Museum of the Academy of Science in Stockholm (1801) and the dry
material, including Queen Louisa Ulrika’s important collection of shells went to Uppsala in 1803. This
specimen may therefore have originated in the royal museum and is possibly part of the type series of 0. cornutus
because Linnaeus based his name in part on the description in Museum Adolphi Fsidnici (Linnaeus, 1754a).
However, the connection is not certain because in Thunberg’s (1787) catalogue of the Uppsala collection a
specimen of 0. cotnutus is listed from the Ahlstrijmer-LinnC donation of 1749, and in the Thunberg manuscript
catalogue of 1828 there are two specimens under this name, the Ahlstromer-LinnC specimen and a second one
labelled beta from a Thunberg donation. As the present specimen has at some time been labelled “beta” this
suggests that it has been identified with the Thunberg collection. Liinnberg (1896) claimed that the
Ahlstromer-LinnC specimen was lost, but no special significance can be placed on this as he appears not have
located dry specimens in the Uppsala collection.
In view of the discrepancy concerning the origin of this specimen it is advisable not to accord it type status.
Tetracdontidae
Sphaeroides tcstudincur (Linnaeus, 1758) [Fig. 281
Tetraodon fcJfUdncus Syst. Nut. (10): 332 (1758); Syst. Nut. 12): 410 (1766).
Specimen: ZIU 47 [39]; S.L. 59 mm, T.L. 75 mm; in spirit, good condition.
LINNAEAN FISHES IN UPPSALA
191
Figure 28. Sphocroides kstudineus (Linnaeus, 1758), (ZIU 47), part of the type series. Above, lateral
view: below, dorsal view.
Label: Tetraodon/testudineus/Mus.Ad. Frid.
Discussion: Linnaeus’s Tetraodon kstudinm was based on two references to earlier literature, Artedi (1738) and
Linnaeus (1749a). No specimens described by Artedi are currently known to exist today. Balk’s dissertation
Museum Adolph Fridcricinnum of 1746 was based on specimens donated by the Crown Prince to Uppsala
University.
The present specimen is listed under this Linnaean name by Thunberg (1787) and in his manuscript
catalogue of 1828, by Unnberg (1896),who recognized it as a type (claiming, incorrectly, that Giinther (1870)
regarded it “as a type”), and by Holm (1957) who also listed it as a type. The curatorial history is thus
complete and there seems to be no doubt that the present specimen is the one described in Balk‘s dissertation
(the illustration, Tab. XIV, figure 3, is a g o d likeness showing the characteristic light bands enclosing circular
and elliptical areas on the back). This specimen is therefore the only type material surviving of 7.lestudineus.
A. WHEELER
192
oceuIJyE (Linnaeus, 1758)
Tetradm wWurSjst. Nut. (10): 333 (1758); Syst. Nut.(12): 411 (1766).
Specimen: ZIU 166 r
e];
S.L.91 mm, T.L. 108 mm; in spirit in fair condition.
Label: TcuaodQnladlatur/Mua. Linn.
Discwskm The taxon Titmdor~o c d b s was based on four earlier accounts, the then unpublished second
volume ofthe M w w n ~ (Lianeetlr,
M
1764b), Osbeck (1757),%nnaeus (1757) and Artedi (1738).
Fernhdm and wheder (1983) have identified a specimen in the National Museum of Natural History,
Stockhoh (NICM 8813) as the specimen described in Linnaeua (1764b), and this is undoubtedly a type
specimen.
There is uncertainty a b u t the type status of the present specimen which originated iq thRAhlstromer-LinnC
ted as such by Thunbag (1787) and in his mankript catalogue of
at it was not conspccifie with Lianaeurand @beck's descriptions,it
Fugu
are a number of species which were described by Osbeck ( 1757) and
onation to the Uppaala A d c x i y . However, this specimen appears to
uncertain date. It i s thus not jwilicd in assuming that it is an Osbeck
specimen, althotlga it may be, but because of this uncertainty it should not be regardad as type material.
It should here be noted pmenthetically that Fernholm & Wheeler (1983) pointed out that there were several
casea where pomibly the same specimens were described in both the M v ~ l n rAd+i
F W (Linnaeus, 1754a
and 1766) aa w d l as by OSbtcL (1757). It is possible that Osbeck specimcna were distributed to both the royal
collection and to Linnaeus.
Arothron AispidyI (Linnaeus, 1758) [Fig. 291
Tetraodmkjdms &st. Nuf. (10): 333 (1758); Syst. Nut. (12): 411 (1766).
Specimen: ZIU 102 [23]; S.L. 93 mm,T.L. 122 nun; in spirit, good condition.
Label: Tetraodon/hkpidus/Mus.Lagerstr. [Note: a second, smaller specimen of S.L. 22 mm, T.L. 27 rnrn in the
same bottk is refern& to Diden.]
Discussion: T"& hispidw was described from two earlier references, the Odhelius dissertation Chinmsia
Lagerstronha (Linnaeus, 1754b), and the work of'Artedi (1738). The present specimen was catalogued by
Thunberg (1787) and manuscript Wng of 1828, by Lijnnbeq (1896) and by Holm (1957) as originating in
the Lageranom donation of 1748 and the latter two recognize it as a type specimen.This seem i d t a b l e and
as no material from Artedi's aemunta airt it is the only surviving type material of T. hispaks.
Diodon hystrix Linnaeus, 1758
Dioda hy& @st. .Ma.(10): 335 (1758); St'. Nut. (12): 413 (1766).
Specimen: ZIU 57
S.L. 262 nun, T.L. 180 mm; dry, stufat, mounted on a wooden base
fw;
Figure 29. Ardtron &+
(Linnaeus, 1758), (ZIU 102). The larger specimen is the holotype of
Tefraodon hupidns (the smaller is an unidentified Diodon sp.).
LINNAEAN FISHES IN UPPSALA
193
Labels: Diodon hystrix Thunb. non LinnC. Chilomycterus reticulatus (L)-this label and redetermination by
Thunberg.
Discussion: The label attached to the base of this specimen seems to have given rise to some uncertainty and
Holm (1957) regarded it a specimen from the “Museum Gustavi Adolphi”, a collection received in 1764. If this
is so it may be the specimen described by Linnaeus (1754a) in the Museum Adolphi Fridericianum which was one
of the four references cited by Linnaeus (1758).
On the other hand a specimen of D . hystrix was listed in the Uppsala collection originating in the 1745
donation of the Crown Prince Adolf Fridrik, by Thunberg (1787) and in his 1828 manuscript collection.
Significantly, the 1787 catalogue entry has a footnote “12) Siccatus” suggesting that this specimen may be it.
Lonnberg (1896) claimed that number 41 “Ostracion conico-oblongus” of Linnaeus ( I 746a) was lost, but
Lonnberg seems to have failed to locate several dry specimens which are known to exist today. Holm (1957)
wrote that the specimen of 7.hystrix from the Crown Prince’s donation of 1746 was missing although present in
1828, but he may have been misled by Lonnberg’s note. As already noted Holm regarded this as part of a later
donation.
Therefore, if this specimen originated from the 1745 Crown Prince’s donation it is probably a type specimen,
but if it came from the 1764 donation of Gustav Adolf it may possibly be a type specimen. On the evidence
available at the present it is impossible to be certain of the history of this specimen.
No other type material of D . hystrix is known.
ACKNOWLEDGEMENTS
My thanks are due in a quite special degree to Dr Lars Wallin, Curator of the
Linnaean collection in the Zoological Institute, University of Uppsala. He has
generously answered all my numerous questions about the Uppsala collection,
and with extreme forbearance tolerated my opening every bottle (but one) of
the Linnaean collection, patiently resealing them after I had finished. His
kindness and generosity as guide to the collection and host in Uppsala made an
essential contribution to this study. I thank Professor Bo Fernholm of the
Swedish Museum of Natural History, Stockholm for hospitality and help during
my visits to Sweden.
At the Linnean Society of London I have to thank Librarians, Miss Gina
Douglas, and Mr Gavin Bridson for access to early Linnaean literature.
Similarly at the Natural History Museum, London, successive Librarians Mr
Rex E. R. Banks, Mr A. P. Harvey and M r M. J. Rowlands provided privileged
access to the Linnaean collection, and the staffs of the Zoology and General
Libraries were everlastingly helpful. My thanks are also due to my former
colleagues in the Department of Zoology, the Natural History Museum,
particularly Mr Patrick Campbell and Mr Oliver Crimmen for much needed
logistic support. I am extremely grateful to Mrs Sita Fonseka who not only
painstakingly typed this paper but brought a degree of organization to it.
This paper was commenced in 1985 but languished due to other
preoccupations in the British Museum (Natural History) that frustrated
constructive work. It was completed in the peaceful surroundings of the Epping
Forest Conservation Centre and I thank M r Paul Moxey, Warden and Director
of Studies, and his staff for the facilities provided there.
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