Carl Linnaeus and the Amphibia

Abstract

Linnaeus did not like snakes. He demonstrated his aversion to snakes in several passages of his writings. He also described toads as “these ugly animals” in a narrative published in 1747. He wrote in the 10th edition of Systema Naturae (1758) in the introduction to the class Amphibia, which he had created already in 1735 and included both amphibians and reptiles: “These most terrible and vile animals are distinguished by their unilocular and single chambered heart, arbitrary lungs, and divided penis” followed by a another long harangue of mostly sarcastic character. In a letter to a friend, when he had just taken a mortgage on a new house in 1758 he uttered: “I have always feared debts as snakes.” In spite of the frowningly spoken approach, Linnaeus dealt with herpetology in systematics and field observations, as well as in his teachings, with the same zealous commitment as he did with the other groups of animals. I have gathered and commented on Linnaeus’s writings on amphibians and reptiles to demonstrate his knowledge and experience in herpetology and how they enhanced over his lifetime. An abbreviated portrayal of his scientific career and biography is imbedded in the text.

Full text

5
INTRODUCTION
In the composition of an overview of
Linnaeus’s contributions to herpetology,
I have chosen to primarily describe his
various achievements chronologically. His in-
volvement in this eld has mainly been based
on his publications, but there are other sourc-
es, such as his correspondence (much of it is
published), his students’ annotations that have
been published and to a certain extent what is
left of collections he worked with. Linnaeus
carried out several journeys in Sweden and
one to Holland that usually resulted in pub-
lished narratives. His diaries during these ex-
peditions have been published posthumously
with annotations. The observations, annota-
tions and contemplations of herpetological
issues in these eld studies hence form an im-
portant part of the present essay. The literature
in English on Linnaeus’s life and contributions
to science as a whole is lavish, e.g. Blunt and
Stearn (1971) and Goerke (1973), two primary
sources that I have used in this essay for his
life history at large, so I provide only a skel-
eton of Linnaeus’s biography, but hopefully
sufcient for any reader to gather the basics of
his life and career.
The Gregorian calendar was introduced in
Sweden on February 17, 1753. The earlier cal-
endar dates were based on the Julian calendar.
I have adjusted the dates used by Linnaeus,
notably for his eld observations, by add-
ing the 11 days difference and marking them
“n.s.” (i.e. “new style”).
Linnaeus did not write anything in English;
he used Latin or Swedish depending on the
avenue of publication. Linnaeus’s own text
or passages that were originally written in
Swedish or Latin are here reproduced in
English. Any text available from previously
translated English sources I have used critical-
ly and sometimes revised, as translators have
not always been observant to the scientic sig-
nicance of precise details.
Linnaeus did not separate reptiles and amphi-
bians into two different classes. Instead he used
Carl Linnaeus and the Amphibia
Richard Wahlgren
Prennegatan 23B, SE-223 53 Lund, Sweden, richard.wahlgren@live.se
Abstract. Linnaeus did not like snakes. He demonstrated his aversion to snakes in several passages of his
writings. He also described toads as “these ugly animals” in a narrative published in 1747. He wrote in the
10th edition of Systema Naturae (1758) in the introduction to the class Amphibia, which he had created
already in 1735 and included both amphibians and reptiles: “These most terrible and vile animals are
distinguished by their unilocular and single chambered heart, arbitrary lungs, and divided penis” followed
by a another long harangue of mostly sarcastic character. In a letter to a friend, when he had just taken
a mortgage on a new house in 1758 he uttered: “I have always feared debts as snakes.” In spite of the
frowningly spoken approach, Linnaeus dealt with herpetology in systematics and eld observations, as well
as in his teachings, with the same zealous commitment as he did with the other groups of animals. I have
gathered and commented on Linnaeus’s writings on amphibians and reptiles to demonstrate his knowledge
and experience in herpetology and how they enhanced over his lifetime. An abbreviated portrayal of his
scientic career and biography is imbedded in the text.
Keywords: Linnaeus, Carl von Linné, herpetology, amphibians, reptiles, biography, Systema Naturae,
dissertations.
© International Society for the History and Bibliography of Herpetology, ISHBH
Bibliotheca Herpetologica, Vol. 9(1-2): x-y, 2011

6
the term “Amphibia” to denote both the am-
phibians and the reptiles and established two
orders: “Serpentia” for the limbless Amphibia
and “Reptilia” for the Amphibia with legs.
He also introduced the 3rd order “Nantes”
for cartilaginous and some other sh, which
I will not deal with in this essay. It was an
antagonist to Linnaeus, Jacob Theodor Klein
(1685-1759), who in 1755 coined the Latin
term Herpetologiae to include any serpentoid
animals (e.g. snakes, amphisbaenians, and
worms; Johnson, J. et al 1984). Herpetology
today embraces reptiles and amphibians.
LINNAEUS’S YOUTH AND
UNIVERSITY LIFE
Carl Linnaeus (Fig. 1) was born on May 23,
1707 (n.s.) in the village Råshult, situated in
the south part of the province of Småland in
southern Sweden. When Carl was two years
old his family moved to a larger house in near-
by Stenbrohult, where his father served as min-
ister from 1709. He was the rst child of ve
and the inspiration for his given name most
likely came from the popular reigning mon-
arch Carl XII (1682-1718). His father adopted
the name Linnaeus from the lime or linden
tree (Tilia cordata), called “lind” in Swedish
and “linn” in the provincial dialect. This is
the standard name employed in English, but
after Carl Linnaeus was ennobled in 1761 he
adopted the name von Linné. A curious mix of
German and French components, this name
was employed retroactively in Swedish (and
occasionally elsewhere) to 1757.
When Linnaeus was seven years old he was
sent to Växjö, 50 km away, for schooling in
preparation for priesthood, as it was a family
tradition and the only occupation his mother
would approve for her oldest son. However,
Linnaeus showed little interest in theologi-
cal subjects, and, probably for that reason, no
brilliance. The teachers recommended to his
father that the boy would probably develop
much better as a craftsman, such as a shoe-
maker or tailor. However, one of the senior
masters at the grammar school and a physi-
cian, Dr. Johan Rothman, had observed the
young man’s interests in botany and natural
history in general. Rothman recommended
that Carl’s parents allow him to study medi-
cine, which was the usual discipline for a natu-
ralist. In August 1727 Carl enrolled at Lund
University. He found a sponsor in his profes-
sor of medicine, Kilian Stobaeus (1690-1742),
who took the new student to his house as a son
with lodging and meals. Linnaeus eventually
got access to Stobaeus’s extensive library and
could accompany his professor on sick calls.
However, the academic tutoring was inad-
equate and he left for Uppsala University after
a year. In Uppsala times were hard, especially
for a poor student, but Linnaeus’s brilliance
was soon unearthed. The dean, Professor Olof
Celsius the Elder (1670-1756), who possessed
a strong interest in botany, provided Linnaeus
with free lodging. From the beginning of 1730
Linnaeus lectured in botany and served as a
very popular guide in the botanical garden. He
tutored the children of Professor Olof Rudbeck
the Younger (1660-1740), famous for his bird
studies, and had access to his rich library. It
was in Uppsala he rst met the slightly old-
er student from the north of Sweden, Petrus
Artedi (1705-1735), who shared his inter-
est in natural history, and had a similar kind
of ingenuity. They even divided the special-
ties among them so that, in a nutshell, Artedi
should focus on the shes and the amphibians
(including reptiles) whereas Linnaeus should
restrict his attention to botany and the other
animals.
LINNAEUS’S TOURS TO THE
PROVINCES OF LAPPLAND
AND DALARNA
Olof Rudbeck had undertaken an exploratory
journey to Lappland (Lapland) in northern
Sweden, an unusual expedition for his time,
which had inspired Linnaeus to plan for a sim-
ilar tour. He applied for money at the Royal
RichaRd WahlgRen

7
Society of Science in Uppsala and pointed out
the opportunity of unusual discoveries and
observations of this remote and unexplored
part of the country. On May 23, 1732 (n.s.),
at the age of 25 he started his ve-months-
long journey on horseback and on foot. He
made careful records for the report to his
sponsor, but also for a separate publication.
However, his diary was not published until
almost a century later as an English transla-
tion (Linnaeus 1811). An annotated Swedish
edition was compiled by Bertil Gullander
(Linnaeus 1969). While the herpetologi-
cal observations are trivial, this diary con-
tains the rst records made by Linnaeus. In
Lycksele he met a woman who complained
that she had three frogs in her stomach (this
was a common kind of belief of the period)
which she claimed to have acquired as frog-
spawn when drinking water earlier in the
spring. Linnaeus recommended tar as a cure
but she had grumbled that she could not re-
tain it. When describing the feeding habits
of the reindeer he declared that they can eat
frogs, snakes and even lemmings, which they
run after so intensely that they can’t nd the
way back to the herd. When he reached the
high mountain areas in Norway, he briey
stated that there are “no eas, snakes and bed-
bugs” there.
An interest in mineralogy emerged in Lapland,
and in December 1733 he made a short jour-
ney to the province of Dalarna (Dalecarlia)
especially to investigate the copper mine at
Falun, then the largest in the world. There
he met the provincial Governor, who would
be the benefactor for another expedition into
Dalarna as far as Norway; that six-week-long
trip was made with a small group of students
during the summer 1734. The narrative was
not published until 1889 (in Swedish). A new
edition with detailed annotations with Arvid
Hj. Uggla as the supervising editor was pub-
lished in 1953 (Linnaeus 1953). An English
edition translated by Andrew Casson came out
in 2007 (Linnaeus 2007a). The company set
off on July 14 (n.s.) from Falun. They stayed
at the Älvdalen parsonage for a few days and
used the time for excursions and observations.
Linnaeus wrote with an annotation in the left
margin “vipera vera” (the “true viper”):
When we returned we found a quarter long [20
cm] snake, its back was with a linea longitude.
Dentate, fusca [a dark line with indentations
lengthwise], besides this the whole snake was
gray apart from blackish spots just opposite
each tag on the entwined line. The belly was
brown with a black spot at the tip of each scale.
Gula [The throat] was pallida [pale] and la-
bium superius, secundum marginem white [the
upper lip white at the edge]. Iris oculorum ig-
nea, cum pupilla perpendicular nigra [The iris
colored as re, the pupil vertical and black].
This was obviously an adder, Vipera berus (L.,
1758).
In Älvdalen they “went out shing and for a
long time tried in vain to observe a 4-footed
sh that was said to run up the trees, one that
we thought to be Salamandra aquatica.” On
August 26 (n.s.), the company on their way
back had reached Floda (now Dala-Floda)
CARL LINNAEUS AND THE AMPHIBIA
FIGURE 1. Portrait of Linnaeus from a hand
colored steel engraving postcard that bears the date
24 May 1907; after a gypsum medallion by C. F.
Inlander in 1873.

8
and Linnaeus reported that carps and “skrot-
abborar” are said to occur there (Sw. abborrar
are perches and the Sw. prex skrot, or skratt,
is probably a dialectal word for a demon). He
commented further on this species within pa-
renthesis: “Salamandra aqvatica or the sh
that is for long said to leap in the trees when
he is on the sh hook.” In local folklore it was
considered to be a sh, but Linnaeus postu-
lated an amphibian afnity even without hav-
ing seen it. Sven Nilsson (1787-1883) in his
classic Scandinavisk herpetologi (1842:108)
assumed it referred to the great crested newt,
Triturus cristatus (Laurenti, 1768), whereas
Ingvar Svanberg (2005) strongly advocated
it to be the smooth newt, Lissotriton vulgaris
(L., 1758), because the dialectical name
is more sympatric with its distribution in
northern Sweden. However, Mattias Sterner
(2005:14) considered it the male of the great
crested newt, which in the breeding season has
a denite appearance of a perch at rst glance
and the actual northern distribution of this spe-
cies is not yet fully investigated. Personally I
consider it even doubtful that the two newts
were differentiated in the local folklore.
Once back in Falun, a well-off man offered
Linnaeus nancial aid to accompany his son,
Claes Sohlberg (1711-1773), to Holland to
study for a few years. This would be a great
opportunity for young Linnaeus to nish his
studies of medicine as well, because due to
a royal decree in 1688 the Swedish univer-
sities stopped bestowing the M.D. degree in
medicine (Goerke 1973:5). During the rst
four decades in the 18th century the univer-
sity in Leiden, Holland, became popular
place for Swedish medical students to study,
and the university at Harderwijk, Holland, to
graduate from. A doctor degree could actu-
ally be acquired promptly and at low cost if
the manuscript to the dissertation was ready
(Holmstedt et al. 1998). It was in 1734 dur-
ing the Christmas celebrations he met Sara
Elisabeth (Lisa) Moraea (1717-1806) to whom
he was engaged in January the following year
and made an agreement with her parents about
a three year absence before they would marry
(Goerke 1973).
HAMBURG AND THE HYDRA
Linnaeus and Claes Sohlberg left for Holland
in early spring 1735, although the pledged
money by the latter’s father was subtly with-
drawn. They made several stops on their way
and stayed in Hamburg in northern Germany
for three weeks. Linnaeus had made himself
known to the intellectual community by hav-
ing his visit announced in a local periodical
dealing with scholarly matters, Hamburgische
Berichte both beforehand and during his stay.
Once there, he sometimes made a show by
dressing in a Lapp outt with the ritual drum
that he had acquired on his Lapland tour and
he also sang his own praises with a collection
of insects plus several scientic manuscripts
that he had plans to publish when he reached
Holland. It was in Hamburg he was shown the
stuffed hydra or seven-headed serpent, which
Albertus Seba (1665-1736), although he had
not seen it himself, had described as a genu-
ine natural specimen and reproduced it from
a sketch in his imposing Thesaurus of natural
history, volume one (1734:158-160, pl. 102)
the year before. The hydra had been looted in
1648 from an altar in Prague and was now for
sale at a substantial price. The body was a foot
long and the tail as much. Linnaeus found that
the skin was taken from snakes, the jaws and
feet from a weasel, and the teeth “were not
naturæ, but artis miraculum.” The hydra has
its roots from the Greek mythology and as a
fabulous animal it has appeared in the natu-
ral history literature, for example in Jonstonus
1653:38-39, plate 11, with reference to Conrad
Gesner (1516-1565). Arvid Hj. Uggla (1946)
has given a detailed story on Linnaeus’s en-
counter with the Hamburg hydra.
From Hamburg Linnaeus and Sohlberg pro-
ceeded down the Elbe on May 28, 1735 (n.s.)
and arrived in Amsterdam after a 16-day voy-
age. Linnaeus wrote in his diary that was com-
RichaRd WahlgRen

9
piled and published by A. Hj. Uggla (Linnaeus
1953; in Swedish):
The frogs croaked very loudly 3 to 4 times
louder than in Sweden, had its own sound.
Some sang so beautifully that you felt new-
born, and banished all disagreeable thoughts,
others so mournfully that one almost dies of
melancholy.
HOLLAND AND SYSTEMA
NATURAE
Linnaeus had already written, in Latin, his
thesis in medicine, “A new hypothesis as to
the cause of intermittent fevers”. Linnaeus’s
theory was that they resulted from living on
a clay soil (Blunt and Stearn 1971). Once
in Harderwijk, it took just a week to have it
printed, defend it in a formal disputation, and
receive the doctor degree, which happened on
June 23, 1735 (n.s.). Within a week, on the
day he returned to Amsterdam, he moved to
Leiden to meet more scholarly people and to
visit the renowned botanical gardens. Among
the many inuential individuals he met was
the botanist and physician Johan Frederik
Gronovius (1690-1762). Linnaeus had many
book-length manuscripts ready, including
the one that would be so fundamental to the
natural science, Systema Naturae (hereafter
SN). Gronovius came from a wealthy fam-
ily of scholars and together with a Scottish
doctor Isaac Lawson (?-1747) he realized its
importance and they offered to pay for the
printing. It was given to the printer in Leiden
on June 30, 1735. SN was in Latin and the
rst book that Linnaeus published (Linnaeus
1735). There are several facsimile editions but
Engel-Leideboer and Engel (1964) have also
provided translation of essential parts. The
‘natural bodies’ were divided into three king-
doms, namely: stones, vegetables and animals.
Linnaeus classied taxa in a hierarchical sys-
tem of classes, orders, genera, and species, a
system still in use although many more levels
have been added. Linnaeus wanted each of the
kingdoms to appear on one page but because
the known animal kingdom at that time was
already heaving, the size of the book had to
be a giant folio (53x42 cm). The size in itself,
and the many tables, caused problems for the
printer in Leiden and it was not nished until
December 9. The book consists of only eleven
printed pages on seven leaves. Each table of
the three kingdoms is preceded by a chapter
“Observationes.” Linnaeus established among
other matters that zoology is the noblest part
of the natural history, that most of what was so
far published on animals consisted of fabulous
stories and imperfect illustrations, and that the
only authors dealing with acceptable system-
atics were Francis Willoughby (1635-1672)
and John Ray (1627-1705). Linnaeus was not
the rst to devise a system for a methodical
arrangement of nature but earlier methods
lacked consistency and were difcult to use
in practice. Nonetheless, the success of SNI
could also be explained by talented marketing
efforts by Linnaeus and his Dutch network.
The table with the orders, genera and species
for the animal kingdom contains six num-
bered classes; these are I Quadrupedia, II
Aves, III Amphibia, IV Pisces, V Insecta and
VI Vermes. Each class occupies one column
in the table, but the Amphibia section is short-
er than the others. Linnaeus identied four
genera, Testudo, Rana, Lacerta, and Anguis,
with altogether 27 “species”, here marked by
me in quotes to indicate his possible concept
of a species in a collective sense, because
he must have realized that there were more
than 27 types of “amphibia” on earth, espe-
cially because he had on June 4 seen Albert
Seba’s extensive zoological collections and
his Thesaurus volumes 1 and 2 (Engel 1936).
Beneath the Amphibia column Linnaeus
noted:
The Creator in his benignity has not wanted to
continue any further the Class of Amphibians;
for, if it should enjoy itself in as many Genera
as the other Classes of Animals, or if those
things were true that the Tetralogists have
fabricated about Dragons, Basilisks, and such
monsters, the human genus would hardly be
CARL LINNAEUS AND THE AMPHIBIA

10
RichaRd WahlgRen
able to inhabit the earth.
[Translation from Latin by Engel-Ledeboer
and Engel 1964.]
The remaining space in the column is lled
with observations on “Paradoxa”, being
mythical animals or phenomena in Nature that
according to Linnaeus lacked natural founda-
tions (Fig. 2). Three of the ten listed paradoxes
have herpetological connotations.
Linnaeus wrote that the seven-headed serpent,
which he had inspected a few weeks earlier,
had similarities with the dragon in the Book of
Revelation and he concluded:
Nature, always invariable, has never in a natu-
ral way shaped several heads on one body.
Fraud and artice, we have easily revealed
from the teeth of the carnivorous weasel in-
stead of the teeth of Amphibians [as dened
in SNI].
It is obvious that Linnaeus had read about the
paradox frog, Pseudis paradoxa (L., 1758), in
Metamorphosibus Insectorum Surinamensium
(1719:71, pl. 71) by Sibylla Merian (1647-
1717) and in Albert Seba’s Thesaurus
(1734:125-126, pl. 78), that the tadpole, which
at a stage is three to four times larger than the
adult frog, develops to a sh:
The Frog-Fish, or the metamorphosis of Rana
into Fish, is very paradoxical, as Nature would
not admit the change of one Genus into anoth-
er one of a different Class.
Linnaeus properly included the ying dragon
from East India as a species, “Draco volans”,
among the Lacerta. To the mystifying “Draco”
that often appeared as a natural curiosity
sometimes fabricated from a ray sh Linnaeus
made these comments:
Draco with a serpent body, two feet and two
wings like a bat is Lacerta alata [=winged liz-
ard] or a Ray articially shaped as monster and
dried.
Linnaeus published in 1736 a broadside with
the title Methodus that explained his method
for the descriptions of natural objects. The
broadside has the same size as SNI and was
apparently furnished with the copies of the
SNI that remained in stock when the Methodus
was printed. Karl P. Schmidt (1952) stressed
the importance of the document, even for
modern practice, for adequate description of
newly discriminated species of plants, ani-
mals, or minerals. He undertook to translate it
for the rst time into English. It contains noth-
ing special about herpetology but it provides a
good representation of Linnaeus’s structured
method of dening names, theory, genus, and
species. (Fig. 3)
In August 1735, on recommendation of
Gronovius and Hermann Boerhaave (1668-
1738), an inuential professor of medicine,
botany and chemistry at Leiden, Linnaeus was
made superintendent of the botanical and zoo-
logical garden in Hartekamp belonging to a
wealthy banker and patron of culture, George
Clifford (1685-1760) (Goerke 1973:28).
Linnaeus spent three years at this country
place between Haarlem and Leiden with some
periods of absence for travels and services for
other people. During this period he had a strik-
ing output of books from manuscripts that he
had produced either in Sweden or in Holland.
Most of those were on botanical systemat-
ics, but he also edited Ichthyologia (Artedi
1738), a treatise on the systematics of shes
from the manuscript of his Swedish friend and
colleague Peter Artedi, who had drowned in
Amsterdam in August 1735.
LINNAEUS IN STOCKHOLM
1738 TO 1741 WORKING AS A
PHYSICIAN
During the summer in 1738 Linnaeus left
Holland for Paris and then sailed directly to
Sweden from France. Linnaeus demonstrated
in letters that he wanted to continue his sci-
entic works as a botanist, but there was no
vacant professorship in Sweden. Instead, he
set up a medical practice in Stockholm, which

11
normally was not a worthwhile business at
the time, but Linnaeus succeeded and with-
in a year he was appointed Physician to the
Admiralty.
In May 1739, Linnaeus participated in found-
ing the Swedish Academy of Sciences which
was based in Stockholm (a discussion of the
early history and contributions to herpetology
of the Royal Swedish Academy of Sciences
has earlier been provided by me (Wahlgren
1999). In June 1739 he married Sara Lisa in
Falun.
Besides serving as a physician he pub-
lished the 2nd edition of Systema Naturae in
Stockholm in 1740 (SNII). The format was
changed thoroughly, produced as a very small
quarto with 84 pages. The text for the class
Amphibia is similar to SNI, but now occupy-
ing one full page. This time he established
two orders: Ordo 1 Reptilia, Pedes quatuor
containing the sequentially numbered genera
79 Testudo, 80 Rana, and 81 Lacerta, and
Ordo 2 Serpentia, Pedes nulli containing the
single genus 82 Angvis. For some species he
added the Swedish vernacular names. He re-
moved the species Rana Carolina and Lacerta
Senembi Mrg. The latter was referred in SNI
to Georgius Marcgravius (1648) Historiae re-
rum naturalium Brasiliae libri octo. In later
editions of SN Linnaeus brought it up again
as a synonym for Lacerta Igvana, the green
iguana, now Iguana iguana (L., 1758). Rana
Carolina never emerged again.
The family’s rst child, Carl, was born in
Falun in January 1741. On May 5, Linnaeus
was appointed professor in botany at Uppsala
University, but it would take about half a year
before the installation and when he gave his
rst lectures.
FIGURE 2. Systema Naturae I – Amphibia and Paradoxa.
Amphibia, comprised of both amphibians and reptiles,
formed Class III of Regnum animale in Carl Linnaeus
Systema naturae 1735. Paradoxa was comprised of
mythical animals or phenomena in nature that according to
Linnaeus lacked natural foundations.
CARL LINNAEUS AND THE AMPHIBIA

12
RichaRd WahlgRen
JOURNEY TO THE BALTIC
ISLANDS ÖLAND AND
GOTLAND
In January 1741 the Swedish parliament in-
vited Linnaeus to undertake a journey to the
Swedish islands Öland and Gotland in the
Baltic Sea. The purpose was primarily eco-
nomic, with goals of identifying natural re-
sources such as new material for dyeing, clay
for porcelain and tobacco pipes, and nding
plants for pharmaceutical purposes, but also
to identify what belongs to historiam natu-
ralem patriæ. Recording of observations was
fundamental and the diary now belongs to the
Linnean Society of London. The narrative was
not published until October 1745 (Linnaeus
1745a). The text is in Swedish, but
Marie Åsberg and William T. Stearn
(Linnaeus 1973; 2007b) have pro-
vided English translations.
He set off from Stockholm in May
26, 1741 (n.s.) on a land route with
a group of six students and ofcers,
and returned on September 8 via
Stenbrohult, the place in Småland
where he spent his childhood and his
father still lived. Linnaeus was in-
deed also a botanist, and he wanted
to study the plants during the ow-
ering season, which is quite short in
Sweden. The group therefore had to
rush through the itinerary in order to
see as much of the blooming ora
as could be possible. On the fourth
day, after leaving Nyköping, they
found a grass snake (Natrix natrix
[L., 1758]) and Linnaeus described
it quite precisely but at this stage he
did not count the scales:
Snakes are neither in Sweden nor
abroad at any point sufciently well
described. Therefore, in order to il-
lustrate historiam naturalem patræ we
will make descriptions of them. Here
we found a Coluber, which hence was
noted: gray body with ovate scales, on its back
more narrow, with a raised linea, in the same
color except a few which are blackish at the tip,
so that his back looked slightly mottled. The
underside was covered with scales as large as
the belly or half of the body. These are black,
lighter at the sides, closer to the head even
more. The head black with several sutures. The
jaws with black transverse stripes. Nostrils
small. Around the ears black with large white
patch. The teeth in two rows on each side and
are of equal size. The tongue is composed of
two black strings. This one makes no harmful
bites and is of the Grass snake genus.
In another ve days the group had reached
Vetlanda in the heart of Småland and they
came upon the second snake, which was the
European adder, Vipera berus (L., 1758):
FIGURE 3. Methodus, a broadside that Linnaeus published
in 1736 in which he explained his method for the descriptions
of natural objects. It has the same size as Systema Naturae I
(1735).

13
An adder lay on the road . . . , he was big and
gray, his back with what resembled a dark
thorny band, between which thorns a blue spot
was observed along the body, the head was
separated from the dark back by a blue an-
gulus acutus, whose bra superior was at the
edge with white spots. In the mouth, on either
side, toward the tip, and in the upper jaw was
a large fang, which could be both retracted and
expanded like a cat’s claw, which revealed that
this snake belongs to the genus that causes
harmful, indeed often fatal wounds.
A month later, on Öland, in a limestone quarry
they found a common toad (Bufo bufo [L.,
1758]) under a stone, which Linnaeus de-
scribed morphologically with accuracy, simi-
lar to the snakes. Just a few days later, on July
1 (n.s.), while waiting for a boat to bring the
group to Gotland they surveyed the area and
could report on the grass snake (Natrix natrix),
which he described morphologically with lit-
tle additions to the previous individual’s char-
acters and a Rana temporaria Charl. onom.
24. The Latin name can be misleading as this
happened before he introduced the binominal
nomenclature.
Rana temporaria Charl. onom. 241 was caught
and described; its hind feet had 6 toes, which
were somewhat webbed, the rst smallest, the
second in order longer, the second last lon-
gest. The fore-feet or hands had 4 toes, well
divided, of which the 2nd and 4th were shorter.
The back was at, separated from the sides by
a raised line, which ran from the point of the
head toward the eyebrows, afterwards to the
tail. She was completely gray with black elon-
gated stripes and warts on the back; the thighs
were even paler with black transverse stripes.
[1Walter Charleton (1619-1707) was an English
writer who published on theology, natural his-
tory, and antiquities.]
Linnaeus eventually listed this individual in the
SNX (1758) under Rana temporaria L., 1758,
the common frog. However, this species does
not occur on Öland or even in the immediate
neighboring mainland. They had either found
a moor frog, Rana arvalis Nilsson, 1841, or an
agile frog, Rana dalmatina Bonaparte, 1840,
but regrettably Linnaeus did not recognize it
as a separate species from Rana temporaria,
as the three taxa are morphologically quite
similar.
The group continued to Gotland but Linnaeus
reported nothing of observed reptiles and am-
phibians on this island. Back on the mainland,
in Orraryd 32 km south of Växjö, on August
13 (n.s.), Linnaeus acted as a physician and
treated a woman, who had been bitten by a
snake. He wrote:
Stung by a snake was a woman, for which I
promptly prescribed olive oil to be taken in
large quantities and this frequently, but a few
days later I heard that this, by the English so
appraised medicament, had no effect what-
soever, although the woman was bitten by a
gray snake (Vipera) only 6 hours earlier, be-
fore she began to take the oil. The Medici and
the Pharmacists then should be concerned to
procure for our pharmacies radicem Senegae
from Virginia.
Linnaeus will later report in more detail about
this North American remedy for the snakebite
(Linnaeus and Kiernander1749) with refer-
ence to John Tennent (who had published a
paper on snakebites in 1738).
When dealing with harmful snakes, Linnaeus
continued the text about the “äsping”:
Äsping, a kind of red snake, said to be small
and thick, but moreover very quick to deliver
deadly bites, was told to be often seen in this
part of the country. Whoever hereafter sees
them does well in describing a non-described
animal; then he should be careful to count the
scales between the chin and the tail on the
underside.
The vernacular name äsping is still used in
some parts of Sweden for small reddish-
brownish adders, Vipera berus (L., 1758),
which usually are females. In 1749 he will de-
scribe it formally from four specimens that he
eventually obtained.
CARL LINNAEUS AND THE AMPHIBIA

14
RichaRd WahlgRen
On August 13 (n.s.) Linnaeus also reported
about a limbless lizard, or slow-worm, Anguis
fragilis, L., 1758:
The slow-worm is a snake so stanch that she
breaks to pieces, as soon as you hit on her.
Her belly is black, the sides purple, the back
grayish with a dark linea, which on both sides
separates the back from the sides. The people
believed that this snake bites only at mid-day,
but the teeth seem to make him innocent.
On their way toward Stockholm, 66 km from
Växjö, they observed another individual.
Linnaeus demonstrated here that he did not
believe in unfounded folklore, but he fully ap-
preciated the value of well-nursed livestock.
A slow-worm (koppar-orm) was seen at the
road similar to the last one (August 2) [August
13 (n.s.)] but with a black spot at the top of the
head. The farm-girls used to pat the snake on
his back with the at hand three times, on the
supposition that the cattle subsequently will
fare well and become well-groomed when they
in turn were patted with the hand of the maids.
The ancient rei rusticæ autores [the authors on
agriculture] knew the same trick, though they
did not prepare the hand with snakes before.
PROFESSOR IN UPPSALA,
THE DISSERTATIONS AND
THE DESCRIPTIONS OF THREE
EXTENSIVE DONATIONS
The Linnaeus family moved in October 1741
to the ofcial residence in Uppsala adjacent to
the academic botanical garden.
During a period from 1744 to 1750, when
Linnaeus had been established as the professor
in botany for some years and being responsible
for the Academic Natural History Museum,
six donations containing zoological speci-
mens were made to the museum. Linnaeus
examined these and for four of them he made
catalogues. Three of the collections not only
contained amphibians and reptiles, but in
fact these animals made up all or the bulk of
them. The catalogues were published in Latin
as academic dissertations and defended by a
student, sometimes called the respondent, who
probably inuenced the contents to some de-
gree, but it is quite clear that Linnaeus is the
leading author. Linnaeus considered the theses
maintained by students under his guidance as
his own work (Smit 1979). Out of the 186 dis-
sertations written during Linnaeus’s profes-
sorship at Uppsala University (1741-1773),
about 30 were devoted to zoological issues.
At least eight of them deal with herpetologi-
cal matters and these theses are discussed in
this paper. The academic procedure associated
with the Linnaean dissertations is provided by
Torbjörn Lindell in the present volume. In the
citations I have here treated the dissertations
as produced by Linnaeus and the respondents
jointly, but with Linnaeus as the rst and lead-
ing author and holder of any conceptions and
opinions.
The three collections are still to a large ex-
tent preserved at the Uppsala University
Museum of Evolution. They have been ex-
amined by several persons after Linnaeus.
Carl Peter Thunberg (1743-1828), successor
of Linnaeus’s professorship from 1784, listed
the materials in order of the several dona-
tions (Thunberg 1787a, b). Einar Lönnberg
(1865-1942) in his catalogue of the Uppsala
University Linnaean collection (1896) identi-
ed the types. Ossian Olofsson (1886-1973)
has left a manuscript “The Linnaean collec-
tions” at the museum in which he reviewed
the type status (Olofsson 1915) of the speci-
mens. Holm (1957) published the most com-
plex account of the collection. Wallin (2001)
compiled the most recent list of the Linnaean
collection of the museum in Uppsala but based
much of the type status on Olofsson and Holm.
Count Carl Gyllenborg (1679-1746), who
was chancellor of Uppsala University, made
the rst donation in two installments, in 1744
and 1745, which contained various preserved
animals or parts of animals, including amphib-
ians and reptiles, as well as plants. A script
copy of the rst inventory of this donation

15
made in 1744 exists in London at the Linnean
Society of London (Holm 1957). Linnaeus and
Barthold Rudolph Hast (1724-1784) as a re-
spondent described the herpetological materi-
al in a dissertation, Amphibia Gyllenborgiana
(1745). Precise descriptions are made of nine
snakes, eight lizards, one crocodilian, four tur-
tles, and two amphibians, altogether 24 spe-
cies numbered sequentially, most originating
from South America and Europe. Linnaeus
explained for the rst time how to distinguish
snakes by counting ventral and subcaudal
scales, because, he claimed, other characteris-
tics such as color and size vary between indi-
viduals while the scale counts are stable. Most
specimens in the collection are still extant and
13 specimens have been assigned type status
according to Wallin (2001).
The Crown Prince Adolf Fredrik made a do-
nation in 1745 of natural curiosities to the
Academy of Uppsala. Adolf Fredrik was
crowned King in 1751. The collection con-
sisted of 87 specimens in ‘bottles’ of Spiritu
vini, predominantly amphibians and reptiles
(Löwegren 1952). The Prince continued to be
a passionate collector of natural history curi-
osities even after this donation. Thunberg has
stated that the donation consisted of duplicates
of his collection (Löwegren 1952). Nothing is
recorded of the provenance of the animals,
but Löwegren assumed that they were pur-
chased in Holland. Linnaeus and the respon-
dent Laurentius (Lars) Balk (1726-1790)
described the collection in Museum Adolpho-
Fridericianum (1746), a thesis that was de-
fended in the spring of that year. Linnaeus
described 64 species that are sequentially
numbered, of which 30 are amphibians and
reptiles, namely, one turtle, three amphibians,
11 lizards, and 15 snakes, coming from both
the old and the new World. Two engravings
accompany the dissertation, one depicting a
Chinese pheasant and the other showing ve
sh species and an anolis lizard with erected
dewlap. Linnaeus referred to this dissertation
as Museum Principis. Wallin (2001) indicated
that approximately twelve surviving speci-
mens have type status.
The animal collection that Linnaeus de-
scribed in the thesis Surinamensia Grilliana
(Linnaeus and Sundius 1748) originated
largely from Dutch Guyana in South America.
The thesis, written in Latin, was defended by
Peter Sundius (1725-1786). In the foreword
is mentioned that the animals were collected
by the plantation owner Pater Gerret with his
son from Surinam and that they were given
to Mr. Claes Grill (1705-1767), who was a
leading Swedish entrepreneur and head of
the Swedish East India Company and had a
great interest in natural sciences. Grill there-
after submitted the collection to the Museum
Upsaliense. Linnaeus described 24 animals
from the collection of which 17, numbered
3-19, comprised Amphibia; No 25 was a
snake from Turkey given by another donor.
Linnaeus emphasized that the Grill collection
contained a new snake genus named Cecilia,
which was described and the specimen nice-
ly depicted in an engraving (168x266 mm).
According to Wallin (2001), the type speci-
men, Caecilia tentaculata L., 1758, is extant
in the museum. The collection also included
a rattlesnake, which was described with the
words “Crotalophorus scutis abdominalibus
CLXXII, scutis caudalibus XXI, paribusque
squamarum III”, which is likely to be Crotalus
durissus (L., 1758). The specimen is now lost
(Wallin 2001). In an appendix he described a
snake that had been donated separately and
came from Turkey (a country then larger than
it is today), which is also depicted in the en-
graving. He will later name it Coluber am-
modytes (Vipera ammodytes [L., 1758]). The
Grill family originated from Italy, where the
name meant “grasshopper.” The inclusion of
the grasshopper “Gryllus thorace…” in the en-
graving was probably a way of honoring the
donor.
THE SWEDISH FAUNA ON
RECORD – FAUNA SVECICA
In 1745 Linnaeus published his book Flora
Svecica (Linnaeus 1745b) that presented the
CARL LINNAEUS AND THE AMPHIBIA

16
Swedish plant life arranged systematically,
and in 1746 he published a catalogue of the
known natural animals occurring in Sweden,
Fauna Svecica (Linnaeus 1746), both written
in Latin. In Fauna Svecica, Linnaeus treated
44 mammals (beginning with Homines – man),
205 birds, 13 reptiles and amphibians, 76 sh-
es (including six cetaceans), 932 insects and
spiders and 87 “Vermes”, comprising mainly
all other invertebrates, a total of 1,357 species
(Nybelin 1946). The class Amphibia appear-
ing on the pages 94-97 and 387 (an appendix)
is divided into the two orders, “Reptilia” and
“Serpentia”. For each species he gave rst
a description of the salient features with a
dozen words. For the snakes, he stated only
the numbers of the abdominal and caudal
scales, a method rst explained in Amphibia
Gyllenborgiana (Linnaeus 1745b). Linnaeus
listed bibliographic references or synonyms
with an index of authors and their publica-
tions at the introduction. Linnaeus must have
disregarded many important references that
indeed are listed under other groups by listing
only three references under Amphibia: Bart.
Rudolph Hast 1745 (Linnaeus and Hast 1745),
Joh. Paulus Wurffbainius 1683 (Wurffbain
1683), and Christ. Franc. Paullinus 1686
(Paullini 1686).
For the rst frog, as an example, “Rana mini-
bus tetradactylis ssis, plantis hexadactylis
palmatis, pollice longiore” (Rana temporaria
L., 1758), Linnaeus listed nine authors and
their publications, including his own “It. œl”
(i.e. the narrative of his journey to Öland in
the Baltic Sea; Linnaeus 1745a). For each spe-
cies, he mentioned habitats and distribution in
Sweden and its vernacular names. A detailed
morphological description in smaller letters
followed. He listed four species of each of the
genera Rana, Lacerta and Anguis (the last be-
ing the snake äsping still wanting), in all 12
species. Linnaeus did not differentiate between
lizards and newts, or between snakes and the
limbless lizard, and he distinguished two spe-
cies of the adder, Vipera berus (L., 1758), and
three of the smooth newt, Lissotriton vulgaris
(L., 1758). Sven Nilsson (1842) in his early
comprehensive account of the Swedish herpe-
tofauna included 17 species. The Swedish fau-
na currently comprises 19 recognized reptiles
and amphibians (Gislén and Kauri 1959). The
agile frog Rana dalmatina Bonaparte, 1840
was found for the rst time in 1907 in Öland;
the pool frog Pelophylax (Rana) lessonae
(Camerano, 1842) was reported in Sweden by
Sten Forselius (1962). Two large engravings
depicting mainly birds are included in Fauna
Svecica. Linnaeus set the symbols ♂ and ♀
respectively for male and female, a Linnaean
invention (Dal 1996). The word “fauna” used
here was also coined by Linnaeus as a match
to “ora”, being derived from the feminine
counterpart to the Roman forest god of an-
tiquity, Faunus. Fauna Svecica was published
with two different title pages, both dated
1746, one in Stockholm, Sweden, and one in
Leiden, the Netherlands. A second edition of
Fauna Svecica was published 15 years later
(Linnaeus 1761).
A COMMISSIONED TOUR TO
THE WESTERN REGIONS OF
SWEDEN
Upon request by the Parliament ve years
earlier, Linnaeus undertook during the sum-
mer of 1746 a study tour to “Wästergötland”
(Västergötland, the western part of southern
Sweden) to document observations on the an-
tiquities, physics, economy, manufacturing,
medicine, customs, and traditions. He was
accompanied by a student as a secretary, Eric
Gustaf Lidbeck (1724-1803), who in 1756
became professor of natural history at Lund
University. They started on June 23 (n.s.)
and returned on August 22. Linnaeus’s narra-
tive, in Swedish, was published the year after
(Linnaeus 1747). On Ålleberg (a 330 meter
mountain plateau near Falköping) they found
frogs and Linnaeus wrote:
Frogs (Faun. 250.) [Fauna Svecica 1746, no.
250] of a large quantity jumped on the rock
RichaRd WahlgRen

17
hill on the north side, which was so steep that
we ourselves not without the greatest dif-
culty and danger could get up; we wondered
all the more how these miserable wretches
could work their way as far up and what they
wanted so high, which otherwise, always seek
the depth.
It was the animal he later would name Rana
temporaria L., 1758, the common frog.
At a shipyard outside Göteborg (Gothenburg)
they found a snake that Linnaeus could deter-
mine to be the harmless Natrix (Latin for wa-
ter snake; Natrix natrix [L., 1758]).
In Uddevalla they found several toads (Bufo
bufo [L., 1758]) close to the stinking plants
hedge woundwort, Stachys sylvatica, and
baneberry, Actaea spicata. Linnaeus was puz-
zled why the toads seemed to have a liking of
odorous herbs. He wrote:
Toads (Faun. 253) [Fauna Svecica 1746, no.
253] reside in shady places, particularly at the
roots of the rock, where this stinking Stachys
489, and the stinking Actæa 431 grow. I do
not know what pleasure these ugly animals
nd in ill-smelling herbs, and I have seen,
how the Toads intruded into the houses, when
Stachys fœtida are brought inside; in Ukraine,
too, where Cotula fœtida 703 is more common
than elsewhere, there are so many Toads that
they occur everywhere near the houses; but as
soon as Cotula vanishes in the countries also
the Toads disappear.
Stachys foetida is probably also S. sylvatica
and Cotula foetida is mayweed chamomile,
Anthemis cotula. The gures in Linnaeus’s
text refer to record numbers in Flora Svecica
(Linnaeus 1745b). He continued:
I set my dog on a large Toad, he bit her, but
trembled and shook his mouth pretty badly
afterwards and it was obvious, that it harmed
him. Nor could he later be persuaded to try
again, so it is possible what LISTER 1 says is
true that if one seizes the animal with forceps,
then she pisses from each wart a white juice,
which damages and poisons.
[1Dr Martin Lister (c. 1638-1712), vice-presi-
dent of the Royal Society and court physician,
is best known as England’s rst arachnologist
and conchologist.]
In Persberg, near Filipstad, they found toadlets
of what Linnaeus thought was a new species,
which he carefully described. They were simi-
lar to what he had described in Fauna Svecica
(Linnaeus 1746) as no. 253, which later would
become Bufo bufo (L., 1758) with reference,
among others, to the Öland journey (Linnaeus
1745a). This species jumped differently and
its forefeet had four toes and its hind feet had
ve. In SNX (Linnaeus 1758) he would name
it Rana Rubeta, a species with no present al-
location but probably is Bufo bufo.
THE SIXTH EDITION OF
SYSTEMA NATURAE
The sixth edition of Systema Naturae, SNVI
(Linnaeus 1748) is actually the third written by
Linnaeus, because SNIII (1740), SNIV (1740),
and SNV (1747) were published in Halle, Paris
and Halle, respectively, by others (Sherborn
1899), although their production was appre-
ciatively acknowledged by Linnaeus. In SNVI
he placed the animals rst, followed by the
plants and minerals, while the arrangement in
previous editions was the opposite. The genera
are numbered sequentially from No. 1 that is
Homo to 241 that is Sertularia (corals) in the
Animal Kingdom with the class III Amphibia
beginning with the genus 86 Cæcilia and
ending with 95 Testudo on the pages 33-37.
Illustrations in a SN appear for the rst time
in the 6th edition; there are eight copper en-
gravings, the third being a composite plate
with one amphisbaenian, three snakes, and
the Draco with upright wings. The number of
forms under each genus has increased consid-
erably from the 2nd edition. The escalation in
number of genera and higher ranks of herpeto-
logical taxa for Linnaeus’s Systema Naturae is
shown in Table 1.
CARL LINNAEUS AND THE AMPHIBIA

18
RichaRd WahlgRen
LINNAEUS’S LECTURES
There are a numerous lecture notes taken by
students who participated in Linnaeus’s teach-
ing at the university. They are archived in
various libraries in Sweden. Einar Lönnberg
(1913) compiled these for the years 1748-
1752. That was the time when Linnaeus just
had released Fauna Svecica (1746) and his
own third edition of Systema Naturae, SNVI
(1748), but he had not yet introduced binary
nomenclature. The section Amphibia inter-
preted by Lönnberg extends from pages 150
through 167, and Lönnberg’s comments from
pages 431 through 446. Amphibia in the lec-
ture notes was arranged as in SNVI with a
chapter for each species, including details
of the typical morphological characters (in
Latin), their toxicity (for snakes and some
lizards), folklore, behavior and other pieces
of information. For some of his observations
of the animals, or what he had gained from
the literature, Linnaeus drew adequate con-
clusions, but in other respects he was still
clinging to various folklore and superstition.
Below follows a small array of the notes from
the students’ hands that appear in Lönnberg’s
account:
Classis tertiary [Class 3] Amphibia.
This is the shortest of all Classes naturales, so
we should praise the all-wise Creator, as this
Genus is the ugliest, cruelest and most poison-
ous, it would have done the other Genera too
much harm if they were many.
The turtle can live headless for eight days; also
frogs live as long though you take from them
the heart, lungs and all that they have inside.
A person can drink without harm snake poison
taken from the blisters.
When someone is bitten by a poisonous snake,
nothing is more important than expeditious
measures. All the serpent bites are not cured
in the same way, but for the most general, we
want to provide cures. The peasants of the
countryside submerge the injured limb in the
soil, which is quite right as the nasty matter is
then extracted, but it is slow. The customary
way is to bind up around the leg or the bitten
place so it swells and the blood may not come
up in the body; then cut around the bitten leg
several punctures and apply the sucking horn
until it is believed that the evil matter and all
the infested blood are removed. Then you slice
a snake or frog and put it on as well as the lard
from the very snake.
Genus 87 Amphisbaena. Occurring in Greece,
Egypt, Brazil and East and West Indies. … The
old saying that there are snakes with two heads
and that they command the body one at a time,
but such monstrous animals with more than
one head nature knows nothing about, nor are
those existing in tota rerum natura.
Genus 89 Coluber, Sp: 6, 7, 17 are all Viperae.
With these are always shown Signum Martis
♂, which means that they have quite sharp
teeth.
Genus 92 Draco. Bontius 1 calls him Dracun-
culus alatus. … He lives in the East Indies in
the trees, but is probably rare and ies only at
midday when the heat is most intense. He is
not larger than an ordinary lizard. … A dragon
exists in Hamburg, which fooled all Curieusa
[Curious observers] in the World, until
Archiater Linnæus came, when he found him
to be articially made and he ranks amongst
the foremost art piece: He has the teeth of
Mustela [weasels] and is made of many snake
skins and has 7 heads made of Mustela. … On
the dragon are maybe as many stories as there
are old women, but all equally false for no
dragon exists in the world than this small one,
having neither treasures to rest on nor re in
the tail for lighting.
[1 Linnaeus (1758) refers to Jacobus Bontius
(1592-1631) who featured “Lacertus vo-
lans” in 1658 Historiae naturalis & medicae
Indiae Orientalis. Elzevirios, Amstelaedami
(Amsterdam), p. 59 (not 57 as given by
Linnaeus in 1758). Bontius was a Dutch phy-
sician and naturalist who resided in Batavia
(now Jakarta) in Java for many years.]
Linnaeus’s lectures in the Uppsala gardens,
at the University, and on the excursions are
legendary. In a letter written in March 1754

19
to his friend the physician Abraham Bäck
(1713-1795) living in Stockholm Linnaeus
(1910:266-267) wrote (in Latin):
Privately I lecture on Amphibians for such a
magnicent audience, the space is barely ad-
equate, and this to the indignation of my envi-
ous friends and resentment of my enemies.
TWO DISSERTATIONS ON
TREATMENT OF EFFECTS
OF SNAKEBITES
In the spring of 1749 Linnaeus was the rst
preses (chairman heading the examination)
and author to two dissertations addressing
herbal antidotes against snakebites that were
discovered in India/Sri Lanka and North
America. Like the other Linnaean disserta-
tions they are written in Latin. My summa-
ries here are from abstracts of the disserta-
tions in the Swedish twice-weekly magazine
Lärda Tidningar (Scholarly Newspapers) that
the industrious publisher Lars Salvius (1706-
1773) started in 1745 in Stockholm. New
publications from Sweden and abroad were
announced and reviewed in the magazine.
Lärda Tidningar is now a scarce publication
but conveniently all reviews of Linnaeus’s
publications as well as short notices relating
to the learned professor were brought together
and reprinted in 2007 by Ove Hagelin, direc-
tor of The Hagströmer Medico-Historical
Library, Stockholm. Although there is no clear
evidence, it is deemed that Linnaeus himself
wrote at least most of the reviews of his own
publications (Hagelin 2007).
The dissertation Lignum colubrinum (Linnaeus
and Darelius 1749) was defended in March by
Johan Anders Darelius (1718-1780). He was
in 1773 raised to nobility and took the name af
Darelli. The opening part in the review of the
dissertation contains the following passages:
In the East Indies there is a snake called Naja
or Cobra de Capello, which is the most ven-
omous of them all. // There is also a little sore
CARL LINNAEUS AND THE AMPHIBIA
1st ed. 1735 2nd ed. 1740 6th ed. 1748 10th ed. 1758 12th ed. 1766
Serpentia Reptilia
1 Draco 1 Draco 2 Draco
10 Lacerta 9 Lacerta 20 Lacerta 43 Lacerta 48 Lacerta
4 Rana 3 Rana 8 Rana 17 Rana 17 Rana
4 Testudo 4 Testudo 3 Testudo 11 Testudo 15 Testudo
Serpentia
1 Amphisbæna 2 Amphisbaena 2 Amphisbaena
9 Angvis 9 Angvis 2 Anguis 12 Anguis 16 Anguis
1 Caecilia 2 Caecilia 2 Caecilia
1 Cenchris 9 Boa 10 Boa
26 Coluber 82 Coluber 98 Coluber 1
2 Crotalophorus 3 Crotalus 5 Crotalus
1 Meantes
27 25 65 182 216
1 One Coluber described in 1771 is included here.
TABLE 1. The orders, genera and number of species of Class III Amphibia (excluding the order Nantes)
that Linnaeus listed in the ve editions of Systema Naturae that he wrote. The genera are here arranged
alphabetically per the spelling in the 6th edition. The total number of species in this class is listed in the
bottom row.

20
RichaRd WahlgRen
marten, known as Quirpele or Mongoose. …
Now, when it happens that this Mongoose en-
counters a Naja, and it is not better than the
Mongoose is wounded by the poisonous ar-
rows of the Najæ, then the Mongoose must die
unless he can get an Antidote; therefore once
he is bitten he runs onto the ground to take any
herb and to seek Antidotes. As soon as he gets
hold of it, he takes courage and again attacks
his enemy, without fear of its venom.
Several authors described and named as
Lignum Colubrinum (snake wood) many dif-
ferent plants that they believed the mongooses
used as antidote. Linnaeus had examined the
characteristics of the various plants with this
name and could identify the correct antidote
as a tree from Ceylon (Sri Lanka).
The next dissertation, Radix senega (Linnaeus
and Kiernander 1749) was defended a few
weeks later by Jonas Andersson Kiernander
(1721-1778). The opening part in the review
of the dissertation in Lärda Tidningar contains
the following passages:
In America exists a common snake, which is
called Rattle snack or [in Swedish] Skaller-
orm, because he has on the tail a rattle, which
the Creator seems to have put there like an
alarm clock to warn people from this snake’s
bite, which is so dreadful, that a person dies
within a few hours while she is coughing blood
and the esh falls off the bones. Yes, the bite
is so venomous that, if the snake in the strike
gets into a large blood vessel, the Patient often
dies within 2, or no more than 5 minutes; this
is before any cure can be applied. The people
of Virginia and in particular in our Swedish
Nation at Philadelphia have with endless
Experiments suffered this effect.
The Native1Americans have, from the arrival
of the Christians in America, had a Powder
made from a root that they used against the
said horrible bites, specically when the snake
did not hit any major artery; which Powder is
always found to be Specic and infallible if it
is applied promptly; even, as the bite has al-
ways been deadly when this antidote was not
available. But the natives were in no way in-
duced to reveal to the Europeans of what kind
of plant the root was, until Rewards were intro-
duced by the British and thus it was discovered
that she is an herb, called Senega.
[1 Swedish: De wilde Americaner…]
It was the Scotsman John Tennent, physician
in Virginia, who became aware it was the root
from a plant Polygala Senega. The disserta-
tion includes an overview of snake venoms
and of the genus Crotalophorus (rattlesnakes)
with its two species, one earlier described in
Surinamenis Grilliana (Linnaeus and Sundius
1748) and the other in Museum Adolpho-
Fridericianum (Linnaeus and Balk 1746).
Torbjörn Lindell provides in the present vol-
ume a more comprehensive account of the two
dissertations Lignum colubrinum and Radix
senega.
A COMMISSIONED JOURNEY
TO THE SOUTHERN
PROVINCE OF SKÅNE
The Swedish Parliament commissioned
Linnaeus in 1748 to carry out a journey
through Skåne (Scania), the southernmost
Swedish province. His task was similar to the
previous expeditions, in short to discover nat-
ural resources that could be used to improve
the Swedish economy. As usual he comment-
ed on many aspects of nature, geology, people,
antiquities, etc. Linnaeus was now 42 years
old and an internationally recognized scientist
in his mid-career. The narrative, authored by
Linnaeus, was published in 1751 in Swedish
(Linnaeus 1751). The diary, which is now in
the Linnean Society of London, was compiled
and annotated by Bertil Gullander (Linnaeus
1975) and it contains valuable additional in-
formation. A few herpetological observations
were reported.
He set off in the second week of May 1749
(n.s.) from Uppsala, this time in a coach to-
gether with a secretary, his student Olof
Söderberg (1728-1758). Linnaeus typically

21
was observant of amphibians and reptiles.
His rst observation was on June 8 (n.s.) at
the east coast of Skåne in Ravlunda where he
commented that snakes are relatively rare, a
somewhat odd remark, because he did not re-
port that snakes should be other than scarce
in any of his narratives. At the same place, he
wrote on June 7 (n.s.):
The frogs sang in the evening, however, they
had another sound than our common Swedish
frogs. They were said to play until midsum-
mer, when ours [common frogs] play as soon
as the ice is gone.
This is most probably the European tree frog,
Hyla arborea (L., 1758), a species restricted
in Sweden to southeastern Skåne. He will later
describe it in SNX (Linnaeus 1758) without
reference to this observation. Presumably he
never collected any specimens. The day after,
in Andrarum, he heard the re-bellied toad,
Bombina bombina (L., 1761):
The farmers call this kind of frog, barley frogs,
because, when they begin to be heard, it is the
right time to sow the barley.
Linnaeus described the re-bellied toad in
Fauna Svecica (Linnaeus 1746), but only with
an obscure reference to Johan Leche (1704-
1764; professor of medicine and bird spe-
cialist in Turku, Finland). Linnaeus did not
include it in SNX, but described it again as
Rana Bombina in the second edition of Fauna
Svecica (Linnaeus 1761) with reference to It.
Scan., thus signifying this journey.
In Skanör, located on the southwestern cape
of Sweden, he found “Lacerta cauda verti-
cillata”, giving reference to “Fn 1352”, i.e.
Fauna Svecica (Linnaeus 1746:387). Linnaeus
conated the two Lacerta lizards in Sweden,
Lacerta agilis L., 1758 and Zootoca (Lacerta)
vivipara (Jacquin, 1787). In the 2nd edition
of Fauna Svecica (Linnaeus 1761:103) he
described the proper Lacerta agilis (α) with
two varieties: β “Lacertus viridis” (being
very rare) and γ “Lacertus dorso punctis al-
bis duplici serie” (observed in Lapland and in
Uppsala); with the latter he meant what is now
Zootoca (Lacerta) vivipara (Nilsson 1842:25-
26). However, in SNXII (1766) he dropped the
γ-variety.
On the return route through the province of
Småland he received news that one speci-
men of the snake Äsping, which he had ad-
vertised for in the travel account published in
1745 (Linnaeus 1745a) and in Fauna Svecica
(Linnaeus 1746), had been caught in Angel-
stad for him and it could be obtained from a
Reverend Ulmgren. Later, although it is un-
clear exactly when, another two were obtained
from a schoolmaster named Kallenberg in
Nötbeck (now most likely Nottebäck). From
the diary it can be understood that Linnaeus
spent some time and effort in obtaining the
specimens, so it is clear that describing this
poorly known snake species was essential to
him. Linnaeus briey portrayed the snake col-
lected by Ulmgren as hissing louder than other
snakes do, that it is ercer, and that the snake
will be described in more detail from the pre-
served specimen.
On August 24 (n.s.) they were back in Uppsala.
The tour had been successfully accomplished
and they had on the whole been lucky with the
weather. This was the last of the longer tours
that Linnaeus embarked on during his lifetime.
A NEW SNAKE DESCRIBED
– ÄSPING
The Swedish Academy of Sciences was found-
ed in 1739 by Linnaeus and ve other individ-
uals, most of them aristocrats. The rst issue
of its quarterly Handlingar (Transactions) was
printed in the third quarter of 1739. Linnaeus
made frequent contributions to the ora
and fauna in the Transactions. The Swedish
language was a nucleus of the Academy,
which therefore was used in all writings and
publications.
CARL LINNAEUS AND THE AMPHIBIA

22
RichaRd WahlgRen
Linnaeus had in earlier years advertised to
obtain individuals of the snake Äsping. On
the journey to Skåne in 1749 he had obtained
three specimens from Småland and he pos-
sessed another, albeit, what he expressed,
doubtful specimen, from Uppsala. On his re-
turn he prepared an account on the Äsping for
the Transactions. In a letter (Linnaeus 1910)
to his friend Abraham Bäck in Stockholm
on October 24, 1749 (n.s.) he lamented that
he was not able to nd an artist in Uppsala
to illustrate the snake. So he must have sent
the preserved specimen to the editor for him
to arrange for the illustration in Stockholm
(although there is no documentation of this
detail). The paper with title Äsping appeared
in the fourth issue of the Transactions of that
year (Linnaeus 1749a). He began the article
pitying the snakes that have no legs to escape
any predator but, at the same time, admiring
the Creator who had invented a system that
science would in due course call mimicry:
Anyone enemy, whom the Creator himself has
fortied, may not be pleasant encountering.
The Creator has on the other hand been so mer-
ciful telling the snakes, not to act in offence;
thus you never observe a snake chasing man,
but only defending himself, when one gets too
close to him.
I halt for admiration, when I carefully reect
on this arrangement by the Creator, in particu-
lar when I notice that the Master of Nature has
not supplied every snake with arms, instead
may often a snake, which is most dangerous,
dress as one without arms, as well as an un-
armed, on the contrary, may carry the armor of
the equipped.
He reiterated what he had said in the disserta-
tion Amphibia Gyllenborgiana:
A few years back, when His Majesty and
Advisor to the King, Count Carl Gyllenborg
bequeathed to me the beautiful collections of
Indian animals, which now Upsala Academy
is proudly in possession of, I received the op-
portunity to examine the snakes; I then learned
that the broad scales on the belly and the small
scales under the tail were just the only marks,
through which the Creator characterized the
snakes. I then arranged all the snakes I had in
the Museo, after this given reason, which can
be seen in Syst. Natur, 6, p. 34.
He described the Äsping in great detail and stat-
ed that it resembles a young adder. Drawings
on plate VI depict the snake from above and
below. Linnaeus nished the description
But getting to the true characteristics, it con-
sists of broad scales that run from the chin to
the tail or the vent, which amount to 150, but
under the tail are 34 divided scales.
When I for the rst time saw this snake I was
in doubt whether it could be a Viper or com-
mon Adder, in particular as the Viper has 145
scales under the belly and 36 under the tail,
which together make 181 scales while the
Aesping’s scales together make 184. But as all
these [four] snakes agree in the minute scales,
the spot on the tail, and other small details in
the scales on the head, I am certain that it is
a true species, which by Naturalists should be
called
COLUBER scutis abdominalibus 150, squa-
mis caudalibus 34.
Its distribution, he continued, is mostly and
predominantly the province of Småland. It
would be some ve years before he would
use the binominal system for the rst time.
Linnaeus eventually called this species
Coluber chersea in SNX (Linnaeus 1758:218;
now Vipera berus [L., 1758]). An abbreviated
English translation of that paper was provided
by Wahlgren (1999).
The very specimen of the four that Linnaeus
sent to the Royal Swedish Academy of
Sciences in Stockholm was kept in their col-
lections, which later were transferred to the
Swedish Museum of Natural History. The
Linnaean type material of the three Vipera
species of Coluber berus, Coluber chersea
and Coluber prester, all currently in the syn-
onymy of Vipera berus, were surveyed by
Krecsák and Wahlgren (2008). The specimen

23
sent to Stockholm and depicted in Linnaeus’s
paper was during the course of the survey
found in the main collection, identied as a
Linnaean specimen and designated the lecto-
type of Coluber chersea, now Vipera berus
(L., 1758).
THE DISSERTATIONS IN A
NEW FORMAT –
AMOENITATES
ACADEMICÆ VOL. 1
Linnaeus published the dissertations collec-
tively as a new edition under his own name
in Amoenitates Academicae (Academic
Amusements), rst published in 1749 (AA1;
Linnaeus 1749b). The rst volume was actu-
ally edited by Petrus Camper (1722-1789) and
was released with no changes to the contents,
but did not include all dissertations. Linnaeus
was not happy with this project and started his
own with all dissertations arranged chrono-
logically. The Camper edition thereby came
to an end. Linnaeus’s edition was entirely re-
set, paginated differently, and sometimes in-
cluded names that had been changed and other
modied data. There are several editions and
volumes of Amoenitates Academicae. Kiger
et al. (1999) provided an historical overview,
listing, and index to the scientic names in
the original Linnaean dissertations. The three
museum collections described as dissertations
were included in AA1 (Linnaeus 1749b) and
it is worthwhile to record the page numbers,
because Linnaeus from now routinely referred
to “Amœn. Acad. I” and not to the original dis-
sertations and often without reference to the
title by name:
V. Amphibia Gyllenborgiana. Respodent B. R.
Hast. 1745; 107-140.
XI. Museum Adolpho-Fridericianum.
Respodent L. Balk. 1746; 277-326, pls xiii-xiv.
[Referred to as Museum Principis or as vari-
ously truncated by Linnaeus.]
XVI. Surinamensia Grilliana. Respodent P.
Sundio. 1748; 483-508, pl. xvii.
ON DISTINGUISHING MARKS
BETWEEN SNAKES –
A CONTRIBUTION IN THE
ROYAL SWEDISH ACADEMY
OF SCIENCES TRANSACTIONS
Linnaeus’s method of diagnosing snakes was
to count the scales under the belly and un-
der the tail. The adder, common in Sweden,
was therefore diagnosed in Fauna Svecica I
(Linnaeus 1746:97) as “Scutis abdominalibus
CXLIV, squamis caudæ XXXIX”[Abdominal
shields 144, tail-scales 39]. A person had sent
in to the Royal Swedish Academy of Sciences
a typical adder found in Stockholm having
153 belly shields and 32 tail-scales. Linnaeus
therefore adjusted his theory (Linnaeus 1752),
and explained
…what is added or taken off of the number of
the belly shields, the same is shortened or in-
creased in equal proportion in the scales under
the tail.
In this case there is a difference of two scales
…which is not of any fundament, as the rst
scale under the chin, the last on the tail, and
the scale by the opening, can hardly be deter-
mined, so that the sum ought to be accurately
the same.
Linnaeus therefore pronounced the snake to be
the common adder and not a new species. We
know today that the sum of the number of ab-
dominal and caudal plates may vary between
individuals, but scale counts are still signi-
cant morphological characterizations of snake
species and sexes. A full English translation
of Linnaeus’s contribution was provided by
Wahlgren (1999).
CARL LINNAEUS AND THE AMPHIBIA

24
RichaRd WahlgRen
MUSEUM S:Æ R:Æ M:TIS
ADOLPHI FRIDERICI REGIS –
A MAGNUM OPUS
King Adolf Fredrik (1710-1771) with his
wife Queen Lovisa Ulrika (1720-1782) occu-
pied the royal throne of Sweden from 1751.
The royal couple followed the fashion of the
aristocracy of those times to keep personal
cabinets of natural and articial curiosities.
The king kept his collection at his palace at
Ulriksdal while the queen kept hers at the pal-
ace at Drottningholm, both located in the vi-
cinity of Stockholm. They invited Linnaeus to
arrange the collections. Linnaeus, starting in
1751 made several visits to the palaces up until
1755 (Lovén 1887). His rst published work
about the collections, Museum S:æ R:æ M:tis
Adolphi Friderici Regis (Linnaeus 1754), dealt
with the animals belonging to the king, most
of which consisted of animals in alcohol, such
as reptiles and sh. Linnaeus focused only on
some animals and did not make a complete
description of the collection. He consistently
used a binominal nomenclature here for the
rst time in a zoological work. The book, a
large folio measuring 51 by 35 cm, was lav-
ishly illustrated with 33 uncolored plates, 23
of them depicting serpentoid animals made
by Olof von Dalin (1708-1763) and Jean Eric
Rehn (1717-1793), and engraved by Jacob
Gillberg (1724-1793) (Löwegren 1952:316).
Of the 318 animals described 90 were of the
orders Reptilia and Serpentia (12 amphibians,
50 snakes, 25 lizards, 2 turtles, and 1 croco-
dile). Except for SNI this is by size the larg-
est single volume that Linnaeus ever produced
(Dal 1996). The text is in Latin and Swedish.
The main part of the collection of the king
was in 1801 transferred to the Royal Swedish
Academy of Sciences. Its collections formed
in turn in 1828 the Swedish Natural History
Museum in Stockholm (NRM). A great part of
the many Linnaean type specimens in NRM
stems from this collection. The French au-
thor, Pierre-Joseph Bonnaterre (1751-1804)
meticulously copied 39 of the snakes from
the engravings in Linnaeus’s work for his
Ophiologie (1790) (Krecsák 2006). The ac-
count of the collection of the queen, all of
non-herpetological naturalia, followed ten
years later (Linnaeus 1764a) in a more modest
volume without illustrations. A second part of
the Adolf Fredrik collection (Linnaeus 1764b)
with 21 herpetological species described (16
snakes, 2 each of lizards and amphibians, and
1 turtle) is bound with it, but with independent
pagination; the pages with the herpetological
part (Amphibia Reptilia, Amphibia Serpentia)
being 35-48.
NATURA PELAGI,
A DISSERTATION ON
OCEANIC ANIMALS
Linnaeus continued his intense publishing
work after the voluminous Museum S:æ R:æ
M:tis Adolphi Friderici Regis I was nished,
but for four years only one title touched on
reptiles or amphibians. This was Natura pe-
lagi (Linnaeus and Hager 1757), a dissertation
with J. H. Hager (17??–1770) as defendant.
The animals in the “deep hole sea” are de-
scribed from the study of the latest travel ac-
counts. The class Amphibia is treated on half
a page only and three species are described.
Testudo Midas (Testudo mydas L., 1758) was
taken from the 1757 narrative of the Linnaean
apostle Pehr Osbeck (1723-1805) but the oth-
er two, Squali and Lophius histrionius, also
referring to Osbeck, are shes in the group
Nantes, an order of sh that Linnaeus arranged
systematically for the rst time in Amphibia in
SNX (Linnaeus 1758).
SYSTEMA NATURAE –
THE TENTH EDITION
Linnaeus had earlier written three editions of
Systema Naturae (1735, 1740, and 1748) and
other authors had provided another six; a list-
ing of the editions was provided by Soulsby

25
(1933). In 1758, Linnaeus published the 10th
edition (SNX), his 4th, which eventually would
mark the starting point of zoological nomen-
clature. The International Code of Zoological
Nomenclature xed its publication date to
January 1, 1758. A second volume was botani-
cal and is not as important because an earlier
botanical treatise, Linnaeus’s (1753) Species
plantarum is considered the starting point for
the nomenclature of higher plants.
In Lärda Tidningar, issue number 13 on
February 13, 1758, there is a detailed essay
on the SNX that Linnaeus probably wrote
himself. Linnaeus was known for habitually
praising himself although this time it is indeed
anonymously:
Mr. Archiatri and Knight published the rst
edition of Systema Naturae in Leiden, 1736
[sic] by which he also won a general standing
and esteem of the learned world. This book has
within 20 years, now for the 10th time been
published.
In the comments on the section Amphibia at
least a touch of his fascination with snakes is
noticeable:
The Amphibians, which include the many
strange and exceptional animals, which gen-
erally represent the largest number in Natural
Curiosity Cabinets are gathered here with their
characteristics marked out, so no one can with
any critical usefulness behold such Cabinets
without this manual. Among these are such a
number of Snakes with distinctive rings and
features, that a reader might well get vertigo
when he considers what abundance the warm
countries suffer from these visitors.
The increase of amphibians and reptile species
from 65 in the 6th edition to now 182 is notable
(Table 1). Kitchell and Dundee (1994) pro-
vided an annotated English translation of the
herpetological section of SNX which is also
available as a pdf on the Internet.
The introduction part of the third class of the
animals contains interesting passages, which
disclose Linnaeus’s true opinion about reptiles
and amphibians, his correct interpretations,
as well as misjudgment of phenomena in na-
ture such as some sh species (Nantes) being
Amphibians.
Class III.
Amphibians.
These most terrible and vile animals are dis-
tinguished by their unilocular and single
chambered heart, arbitrary lungs, and divided
penis.
Most amphibians are rough, with a cold body,
a ghastly color, cartilaginous skeleton, foul
skin, erce face, a meditative gaze, a foul
odor, a harsh call, a squalid habitat, and ter-
rible venom. Their Author has not, therefore,
done much boasting on their account.
A polymorphous nature has bestowed a double
life on most of these Amphibians: granting that
some undergo Metamorphosis and others cast
off their old age. Some are born from Eggs,
whereas others bear naked young. Some live
variously in dry or wet, whereas others hiber-
nate half the year. Some overcome their prey
with effort and cunning, whereas others lure
the same prey to their jaws as if by magic.
REPTILES. Footed and have at-nude ears
without ear lobes. They pursue various lives
depending on their structure. Testudines [the
turtles] are protected by their shell. Dracones
[the gliding lizards] y on Wings, whereas
Lacertæ [the lizards] ee on feet, and Ranæ
[the frogs] are hidden by Location. Nor do
they all lack Venom, for example the Toad,
Salamander, and Gecko.
SERPENTES. Footless and, lacking Ears,
are deaf. Lungs separate them from the Fish,
as do Eggs in a chain and a divided penis. In
short, the resemblance of the serpents with the
lizards and that of the lizards with the frogs
is so great as to admit no boundaries. Nature
the savior has armed these creatures, cast onto
the bare ground, ignorant of the use of limbs,
and exposed to every harm, with weaponry
bristling with dreadful venom, each unto its
own kind. These Weapons are very like teeth,
CARL LINNAEUS AND THE AMPHIBIA

26
RichaRd WahlgRen
but they are located on the outside edge of the
upper jaw and can be extended and retracted
at will. They are equipped with a sack of poi-
son which they inject into the blood through a
wound-- the cause of dire results though in oth-
er respects it is inert. And thus these Catonians
have a poisonous bite and threaten death with
the tooth; the cups lack death surely accord-
ing to Redi. He who was in charge armed (♂)
only a tenth of the species, but lest those who
were deprived of the weapons the others pos-
sessed should be miserable and rage too much,
he wished them to be similar in shape so that
all of them, of dubious identication, would be
feared by all. But man’s Benefactor gave to the
people of India the Mongoose along with the
Ophiorhiza, to the Americans the Pig along
with Senega and to the Europeans the Stork
along with the Olive.
Should one wish a diagnosis for these, let him
take it from the presence or absence of feet and
from abdominal and caudal Scutes. But lest
the number, taken from one and added to an-
other, should confuse, it is useful to have each
one numbered (Act. Stockh. 1752, p. 206.). The
Length should be given to and from the anus
and in some cases it should be by color. Be
careful, however, lest the tail, once cut off, has
been regenerated.
NANTES the aquatic nned ones
(Chondropterygios, or the so-called cartilagi-
nous Fishes). A class of amphibians that have
arbitrary Lungs, although it is true that they are
not to be seen [the lungs are pectinate, nned
like those of sh but are joined to an arcate,
cylindrical, bulbous passageway, lacking a
bony rod, unlike that of a sh, except in ex-
ternal appearance]. They do not breathe with
free, but with joined gills. The males lie upon
the females with a divided penis! The eggs
are in a chain with young, the skin is foul, the
bones and the rest are cartilaginous. Nor are
they entirely unschooled in Venom, as witness
Pastinaca [the sting ray] and Torpedine [the
electric ray].
AMPHIBIOLOGI are the smallest of them all,
but none are true. Seba has collected and delin-
eated a tremendous number of them unknown
to himself, but he multiplied them and de-
scribed them but minimally. Catesby sketched
a few serpents more beautifully than he made
notes about them. [Translation from Latin by
Kitchell and Dundee 1994; italics, name of
genera and use of initial capital letters here ad-
justed after the original text.]
A MONOGRAPH ON SNAKES
AND SNAKE BITES – DE
MORSURA SERPENTUM
De morsura serpentum (Linnaeus and Acrel
1762) (Fig. 4), “On Snakebites”, is one of
Linnaeus’s few academic dissertations de-
voted entirely to zoology although technically
it is a medical treatise, Disputatio medica. It
solely deals with snakes and their toxins. It
was defended under Linnaeus’s presidium on
June 16, 1762 and the respondent was Johan
Gustav Acrel (1741-1801), who became a
physician and from 1788 was professor in
Stockholm.
FIGURE 4. The title page of the dissertation Morsura
serpentum 1762 by Carl Linnaeus (preses) and Johan
Gustav Acrel (respondent).

27
The slim booklet is in small quarto with 20
pages. Some of the copies include a rather
tiny illustration, about 60 mm in height and
70 mm in width, engraved and attractively
framed with an ornament (Fig. 5). It depicts
the skull of a solenoglyph snake and has nely
engraved gure labels ‘a’ to ‘h’ with captions
printed further down the page. It is named
Vipera vulgaris, but Linnaeus has no such
name in SNX. It translates “Common viper”
and could indicate that Linnaeus didn’t know
the origin. However, Fredric Hasselquist
(1722-1752), a Linnaean apostle who trav-
eled in the Middle East had, at his premature
death in Smyrna (now Izmir), his manuscript
ready for the narrative in which he not only
described his journeys but also included an
extensive zoological and botanical systemat-
ic part (see Adler p. X in the present volume).
His collections and manuscripts were bought
by Lovisa Ulrika, the Queen of Sweden, and
the manuscript was compiled and published
by Linnaeus in 1757. Hasselquist told how
he on July 3, 1750 purchased from a female
snake charmer “4 different kinds [of snakes];
these I described and preserved in aqva vitæ.
These are Vipera vulgaris, …” The name is
not repeated in Hassselquist’s systematic part,
but still not being binominal, the authors de-
scribed the same snake as Coluber (Vipera)
scutis abdominalibus 118, squamis caudalibus
40, cauda aculeate. In SNX it became Coluber
Vipera (Cerastes vipera [L., 1758]). Thus, it is
possible that the species from which the skull
was taken is a Cerastes vipera.
Svenska Linnésällskapet (The Swedish
Linnaean Society), established in 1917, started
in 1921 the publication of a series with transla-
tions of the Linnaean dissertations from Latin
to Swedish. So far 78 accounts have been pub-
lished. De Morsura Serpentum, no. 47, was
published in 1965 with a reissue in 2006. The
translation was made by Ejnar Haglund (1905-
1991) with the notes and conclusion written
by Telemak Fredbärj (1895-1975). The the-
sis was included in Amoenitates Acadamicæ
volume 6 (Linnaeus 1763) and translated into
English (Linnaeus 1781).
The thesis is divided into seven chapters each
with a headline.
Introduction
Linnaeus begins with the observation that any-
thing taken in excessive quantities can be con-
sidered a poison, but also makes a more strict
characterization of poison as a substance that
when it is applied to the human body internal-
ly or externally, even in small doses, produces
dramatic effects, which are disastrous for the
health and life. He then examines the origins
of toxins that appear in each of nature’s three
kingdoms. In the mineral kingdom there are
arsenic and other metal compounds, in the
plant kingdom a number of toxic genera and
species and in the animal kingdom snakes,
frogs and toads, and “all other concealed.”
Amphibia
The chapter begins with a description of the
amphibian class, which includes disapproving
wording:
FIGURE 5. Only some copies of Morsura serpentum
1762 include an engraving of a snake skull that
Linnaeus titled Viperæ vulgaris. The captions to the
illustration appear in all copies of the thesis.
CARL LINNAEUS AND THE AMPHIBIA

28
For almost all of mankind as well as for other
animate beings the serpents appear, because of
their poison and other unexplained features,
which would thwart our ingenuity, so horric
that they instill fear even in the most daring, if
they unexpectedly slither towards him. … In
the amphibian class animals occur, equipped
with more than peculiar features, which do
not exist in other animals. Most of them live
both on land and in water, and most spend
half the year in death-like winter dormancy.
Their breathing is very different from birds
and mammals, for they inhale the air without
a corresponding exhalation, at least not a no-
ticeable one. Their body is by nature rigid and
abounding in cold blood. They are equipped
with double penis, as far as I know does not
exist in other animals, other than possible in
some insects. With its hissing sound, terrible
countenance and its stinking smell they offend
our senses. Some of them lay eggs, others give
birth to live young, and like the craysh to the
result of the cartilaginous skeleton each spring
take off their old skins, whereby no specic
limit is set for their growth and size. …
The author then discusses the prey capture
methods by snakes with an example from a
Boas Javanensis that swallowed a whole buf-
falo. About the native adder, Vipera berus (L.,
1758), he writes:
Our Colubri Beri also occur with expanded
bodies and seen through dissection they had
swallowed a frog or a mouse or, as we ob-
served, 7 still undigested nestlings.
Tela Serpentum
(The Weapons of the Snakes)
“No scientic issue has been the subject to
more different opinions than that of snake
venom” Linnaeus begins. He tells that some
believe that the venom comes from the tail;
others think it comes from the black forked
tongue but most have the erroneous opinion
that the harm derives from the very fangs, but
that all these theories are monstrous fabrica-
tions and contradicting all careful observa-
tions. About the jaw apparatus he says:
Inside the poisonous snake’s lips sits at the
tip of the upper jaw a little bone, which the
snake can move back and forth. This bone is
attached to two or three fangs, larger than all
other teeth. The snake can move the fangs with
help of the little bone similar to the cat with
its claws: erect them when he is provoked and
hide them when he goes to rest after the bite.
These fangs, which common folk call Dentes
majores serpentum have been valiantly de-
scribed by Tyson. Act. Angl. p. 144 in Anatome
Crotali [Edward Tyson (1683) Vipera caudi-
sona Americana, or The Anatomy of a Rattle-
Snake. Philos. Transact. (Royal Soc.), No.
144.]: …Each fang is surrounded by a small
bladder, equipped with glands, so that at a
pressure on the glands a uid seeps out and
can be seen from the very tip of the fang.
Virus (Poison)
In the past it was thought that anger is inamed
from the dark bile, a view that probably lacks
all likelihood, but nevertheless without hesita-
tion it is invoked in this context. For, by virtue
of this, by the Gods uncertain hypothesis, the
origin of the snake’s venom has been deduced.
Namely, it was envisioned that some unknown
duct, which would directly lead the bile from
the gallbladder to the snake’s mouth, from
which the bite would oat into the wound, and
then the deadly symptoms were caused. But
let us leave the ancient delusions and glance
at the far truer observations, gained from more
recent time earnest naturalists.
Linnaeus gives references to Francesco Redi
(1626-1697) who conducted several experi-
ments with vipers and their poison. Linnaeus
relates the divergent views between Redi and
Moys Charas (1619-1698) and the debate a
century earlier (see Knoefel 1988). Linnaeus
dismisses Charas’s opinion that the poison
is generated by a are-up of natural spirits,
and thus snakebites were not fatal, unless the
snake had been teased. English surgeons have,
Linnaeus says, injected medicine diluted in al-
cohol in the blood, with some medical success,
but that the patients usually died. Linnaeus
does a matching with snake venom and milk,
man’s rst nourishment, which also leads to
death when applied in the veins. In the same
RichaRd WahlgRen

29
way as a little yeast can ferment large dough,
the entire blood mass can be destroyed by
snake venom injected to the blood, but how,
Linnaeus dares not say.
Puncturæ symptomata
([Snake] bite symptoms)
In this chapter, the author aims to account
for the various genera and species and the
different toxins. Linnaeus rst depicts the
typical symptoms following the bite of the
native adders, Vipera berus (L., 1758), which
he refers to as “BERI nostri”, our Beri. He
then describes the snakes found in the clas-
sical literature: “HÆMORRHOUS, DIPSAS,
and ASPIS, the latter of three different types
Phtyas, Chersea, and Chelidonia.” Linnaeus
refers to the results of the tests conducted by
Cleopatra’s physicians on persons sentenced
to death. Aspis produced the gentlest death
and this is according to Linnaeus probably the
serpent that the queen used for her demise.
He refers to SNX (1758) in which the poison-
ous snakes, “in addition to if not all the rattle-
snakes”, are listed (under the genus Coluber):
Vipera, Berus, Lebertinus, Naja, Dipsas,
Atropos, Chersea, Severus, Atrox, Mysterizans,
Leberis, Prester, Stolatus, Niveus, Ammodytes,
Aspis, Lacteus, Corallinus. Actually, Linnaeus
did not list in SNX “Prester”, i.e. Coluber pre-
ster L., 1761 (Vipera berus [L., 1758]), but
all the others were indeed labeled poisonous
along with three rattlesnakes. Of these, most
now belong in the family Viperidae, two in
Elapidae (Naja naja and N. nivea), ve in
Colubridae, and two have not been allocated
(C. Leberis and C. Dipsas). He particularly
mentions Naja that can attack people with a
bite so intense that the victims die in less than
an hour with symptoms of esh that falls off
from the bones and dissolves to a sticky stink-
ing mass.
Medela (Cures)
Linnaeus explains that the foremost among
treatments is to remove as soon as possible
the poison from the affected area. This can be
done most easily by sucking, but the person
that sucks must have a gum that is not attacked
by scurvy. One can with a knife broaden the
wound and try to draw the poison with the help
of a pumpkin (Cucurbita). Olive oil is particu-
larly used in Europe; the patient should drink
a large quantity. Linnaeus, however, shows a
hesitation for its healing powers. The antidote,
which is mainly used in America, is powdered
“Radix Senegæ”, as well as other American
drugs, such as Eryngium fœtidum, aquaticum,
etc. The author, however, chooses to ignore
them because our knowledge of them is less
certain. With regard to Asia, it is Ophiorrhiza
mungos, which is the famous antidote against
the “colubri Najæ”. The use of Strychnos col-
ubrini appears to be, if not completely worth-
less, of uncertain effect.
The author concludes the chapter by mention-
ing the snake stone, “Lapis Serpentum, Pedra
de Cobras di capello” and states that Redi has
made many attempts to show any effects with
it without success: “Many have told me that
the stone is articial and does not consist of
anything else than burnt deer antler, Cornu
cerviustum.”
Excantatio ([Snake] charming)
Linnaeus concludes the thesis with a short
chapter that deals with snake charming. He
says that according to Ovidius, Plinius and
other authors from ancient times, the Psylli
in Libya could handle the most venomous
snakes and they treated snakebites by suck-
ing out the venom. Linnaeus refers to Fredric
Hasselquist (1757:70) who has witnessed
the snake charmers’ art but that the charmers
did not want to reveal the magic of their tal-
ent. The dissertation nishes with the preses,
Linnaeus, appearing in third person but with a
hope expressed jointly:
Mr. JACQUIN [Nic. Jos. von Jacquin (1727-
1817)], who returned from West India, said in
a letter to Mr. PRÆSES [Linnaeus] that he has
acquired the art of snake charming for gold.
CARL LINNAEUS AND THE AMPHIBIA

30
RichaRd WahlgRen
If this means chewing his Aristolochia angui-
cida, or any other method, is so far unknown
to us but we hope that Mr. Jacquin is about to
publish this secret, which all inquisitives along
with us beg for.
NEW AMPHIBIAN GENUS AND
SPECIES DESCRIBED – SIREN
LACERTINA
Linnaeus often told in letters about his cur-
rent projects to friends and colleagues. The
available correspondence is accessible on
the Internet under the title The Linnaean
Correspondence with summaries in English
and a few with complete translations and
some with scans of the originals (http://lin-
naeus.c18.net). From these letters or summa-
ries I have been able to record the background
to Linnaeus’s description of the greater si-
ren. It all started in a correspondence by let-
ters in the mid-1750s with Alexander Garden
(1730-1791), a Scottish/American doctor of
medicine living in South Carolina in North
America. Garden sent various collected speci-
mens of animals and plants to Linnaeus. He
enclosed in a letter dated May 18, 1765, a
small specimen of an animal and wrote (origi-
nal in Latin):
...I have sent you the characters of a very ex-
traordinary animal, which I have never seen
till lately, though they are common here. I
have likewise sent to you two specimens of it,
though they are much less than the one from
which I made out the characters, which was
between two and three feet in length. I have
sent it to Mr. Ellis for the museum. It happens
to be a middle link between the Lacerta and
Muræna, having some things in common with
both, and yet differing from both. [Alexander
Garden to Carl Linnaeus, 18 May 1765, The
Linnaean correspondence, linnaeus.c18.net,
letter L3591 (consulted 18 October 2010).
Also in Smith (1821).]
Linnaeus responded on December 27, 1765,
saying about the specimen that it seemed to be
a larva of a lizard, if it is not a very special ge-
nus that could be named Sirene [Carl Linnaeus
to Alexander Garden, 27 December 1765, The
Linnaean correspondence, linnaeus.c18.net,
letter L3682 (consulted 18 October 2010)]. He
could not determine if it was a larva or a full-
grown individual. However, it had two feet
with nails, and it could produce a sound. In
a letter to his close friend Abraham Bäck on
January 1, 1766 Linnaeus wrote (original in
Swedish):
A Siren has been sent to me from America;
but a species that is small … she has only 2
arms and hands; no feet. the rear part is like
an eel. She has both gills (Branchias) like sh,
and lungs 2 like the perfect animals. [Carl
Linnaeus to Abraham Bäck, 1 January 1766,
The Linnaean correspondence, linnaeus.c18.
net, letter L3704 (consulted 18 October 2010).]
Linnaeus wrote on January 24, 1766, to Johan
Andreas Murray (1740-1791), a Swedish
professor of medicine and botany living in
Göttingen, Germany (original in Swedish):
Now I amuse myself with an animal that is like
a Siren and has come from America. has only
2 front feet with arms and hands, no hind feet,
but looks like an eel. has both lungs and bran-
chias. sings like a bird. I plan to write about
this. [Carl Linnaeus to Johan Andreas Murray,
24 January 1766, The Linnaean correspon-
dence, linnaeus.c18.net, letter L3518 (consult-
ed 18 October 2010).]
In another letter to Bäck on February 11, 1766,
he wrote:
Now our draftsman is working on the illustra-
tion of the Siren. … The dissertation I am done
with. [Carl Linnaeus to Abraham Bäck, 11
February 1766, The Linnaean correspondence,
linnaeus.c18.net, letter L3719 (consulted 18
October 2010).]
The dissertation (Linnaeus and Österdam 1766)
was printed for hearing on June 21, 1766, with
Linnaeus in presidium and Abraham Österdam
(1745-1776) in defense (Fig. 6). Österdam at-
tended Uppsala University in 1764-1766 and
was appointed Royal Life Surgeon in 1775.

31
The dissertation was reviewed in Swedish in
Lärda Tidningar, No. 5:203-204 (see Hagelin
2007), probably written by Linnaeus himself.
The review has provided me with some keys
to the contents of the dissertation which is in
Latin. The rst two chapters deal, according
to the review, with the mythical Siren, the
third on the differences between the orders
in the Animal kingdom, the forth on strange
animals of our time, and the fth on charac-
ters of shes. The description of the siren is
made in the sixth chapter with reference to
Dr. Garden. This text is a translation from the
review:
In §. 7. is shown to us how the animal di-
verts from the general characterizations in the
Animal kingdom; in lack of hair and feathers
it can be neither Mammal nor Bird; among
Amphibia it cannot be assigned to any of the
known divisions, neither to the Reptilia, which
have four feet and no gills; nor to the Snakes,
that are without feet, ns and gills; nor to
Nantes, which all have ns. In view of this,
Author shows the necessity of establishing a
new Ordo in Reptilia, with two feet, lungs and
gills. Many would think this animal is simply
a larva of Lacerta Iguana or another, but both
claws and its sound deny this, which can be
seen of §. 8. In the end, the animal is described
as Siren Lacertina, of which our Auctor has
done a beautiful copper engraving.
A copper engraving accompanies the disser-
tation (Fig. 7) in which the siren is depicted
with a mermaid named Siren Bartholini that
a Danish author Thomas Bartholin in 1664
had illustrated based on a hand and a rib sent
from Brazil and then called it Homo marinus
(Broberg 1975). The conclusion part of the text
in the dissertation is modied in Amoenitates
Acadamicæ, volume 7 (Linnaeus 1769) and
with a new and larger illustration of Siren lac-
ertina. A systematic part is listing the siren as
in SNXII. The caption to the illustration says
that “Siren Bartholini” is “forte cta” (possi-
bly it is ction).
The greater siren (Siren lacertina L., 1766) is
a valid species; Linnaeus and Österdam jointly
should be the authors of the description and
not Österdam alone, as have been suggested
by Dubois (1989-1990). The type specimen is
extant in the Uppsala University Museum of
Evolution.
SYSTEMA NATURAE – THE
TWELFTH EDITION
Systema Naturae in the 12th edition, SNXII,
with an appendix in Mantissa plantarum al-
tera (Linnaeus 1766, 1767, 1771) was com-
piled by Linnaeus; this being the last of the
ve editions of SN that he was personally re-
sponsible for. It is a much-expanded version of
the SNX edition, although the increase in num-
ber of Amphibia species is less remarkable.
The numbers of herpetological taxa in all ve
editions are shown in Table 1. Linnaeus actu-
ally dropped three species from SNX: Rana
CARL LINNAEUS AND THE AMPHIBIA
FIGURE 6. The title page of the dissertation Siren
lacertina 1766 by Carl Linnaeus (preses) and
Abraham Österdam (respondent).

32
variegata, Rana Hyla, and Lacerta hispida
without leaving what I can nd any reason for
the omissions. In both SNX and SNXII there is
for Amphibia a table of all genera preceding
the listings, which are named Characters am-
phibiorum in SNX and Generum Characteres
in SNXII. He changed within the order Reptilia
the arrangement of numbering and listing of
the genera. These are in SNX: 103 Testudo,
104 Draco, 105 Lacerta, and 106 Rana. In
SNXII they are: 119 Testudo, 120 Rana, 121
Draco, and 122 Lacerta. The number of the
Genera follows a series that was expanded in
SNXII, but I cannot see any explanation for
the move of Rana.
The Siren he placed in a new order Meantes
with the single genus and species Siren
Lacertina. It is included in an Addendum to
part 2 of the rst volume that was published
in 1767. In the Mantissa plantarum altera
appendix (1771) Linnaeus included Coluber
crotalinus, a snake in “Museo Regio” with no
current allocation.
DELICIAE NATURAE
In December 1772 Linnaeus held his last ora-
tion, Deliciae naturae (Nature’s delights) to
mark his departure from the presidency of
the University. It was a brilliant oration held
in Latin by the now elderly Linnaeus. His
students required a translation into Swedish
(Linnaeus 1773). In one chapter of the oration
he compared the animals with an infantry,
the mammals are the foot soldiers clothed in
furs, the birds the cavalry beautifully dressed
in dyed down. Linnaeus demonstrated that
he retained his aversion against the animals
of the class Amphibia. They constituted “an
unsightly, hideous naked mob, with no uni-
forms, inadequately armed except some who
got terrifying poisoned darts. They stick to
the ground; a few are seamen.” The shes
and insects, in turn, he described generally
in positive terms. However, Vermes are also
quite depressingly portrayed as small, slug-
gish mariners with no attire.
THE LAST YEARS
Linnaeus was now an old man. He had his rst
stroke in 1774, when he was lecturing to pri-
vate pupils, and the second in 1776. Short pe-
riods of physical revitalization did not prevent
the trend of gradually failing health. In 1775
he was still teaching students, though more
often than not they could hardly understand a
word he said. On 22 June 1775 he presided,
for what proved to be the last time, at a dispu-
tation. Linnaeus died on 10 January 1778 at
eight o’clock in the morning in his home in
RichaRd WahlgRen
FIGURE 7. The illustration of Siren lacertina,
Linnaeus & Österdam 1766. Linnaeus received the
specimen from Dr. Garden in North Carolina in North
America. Linnaeus could see similarities with the
mermaid that Thomas Bartholin had reported in 1664
based on a hand and a rib sent from Brazil.

33
Uppsala. The funeral took place on 22 January
at the Uppsala Cathedral, where he is buried
(Blunt and Stearn 1971). Linnaeus’s library
and specimens went to his son Carl, now hav-
ing the professorship after his father. However,
after his premature death in 1783 they were
returned to Linnaeus’s widow. She in turn sold
all books, specimens and correspondence the
following year privately to a James Edward
Smith in England. Eventually all ended up in
The Linnean Society of London, where they
still remain except for the mineral collection
that had been sold (Blunt and Stearn 1971).
However, Linnaean zoological materials that
belonged to the Academy of Uppsala, such as
the donations catalogued by Linnaeus were
not included in the sale and largely remain in
the museum.
ACKNOWLEDGMENTS
Torbjörn Lindell has been an enduring partner
for discussing problematic Linnaean issues
and proposing academic sources. Lindell also
read and commented on a draft of the paper, as
did two anonymous reviewers. Lars Fröberg
at the Botanical Museum of Lund University
assisted in identifying pre-Linnaean plants.
Ralph Tramontano assisted with the English
style and grammar. Last but not least I thank
the Herpetologists’ League for support of the
Linnaean Tercentenary Symposium at the Joint
Meeting of Ichthyologists and Herpetologists
in St. Louis, Missouri.
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