Reid • Linnaeus’ life and relationship to modern biology TAXON 58 (1) • February 2009: 18–31

Carolus Linnaeus (1707–1778): his life, philosophy and science and its
relationship to modern biology and medicine
Gordon McGregor Reid

Director General, North of England Zoological Society, Zoological Gardens, Upton, Chester CH2 1LH,
U.K. g.reid@chesterzoo.org

The system of binomial nomenclature and wider taxonomic paradigm forged by Carolus Linnaeus in the
18th century came from his original approach to understanding the natural world. It was also a product of
environmental, economic, social, cultural, political and theological influences of the time. For Linnaeus the
identification, naming and classification of different kinds of animals, plants, diseases, fossils and rocks had
practical as well as theoretical importance. In his life and work he clearly demonstrates the ‘scientific ap-
proach’ including careful information gathering, exploration, empiricism, dissection, accurate observations and
published descriptions. There is an inspirational use of morphological characters in comparative diagnoses, a
requirement for material evidence to support hypotheses and the systematic and hierarchical organisation of
knowledge. However, techniques for specimen preservation and analysis were limited and sample sizes too
small to properly characterise wild populations. Thus he bequeathed artificial and ‘typological’ more than
biological concepts determined by form and pattern rather than process. Species and genera were regarded as
fixed, objective entities. This is sometimes associated with the supposedly stultifying effects of Aristotelian
essentialism on Linnaean and later taxonomy. ‘Natural’ classification, as far as developed, was not phyletic
but, instead, reflected Wolffianism, a modified creationist doctrine. Nonetheless, Linnaeus was the first to
formally recognise the close affinity between humans and primates, a controversial idea later fully developed
by Charles Darwin. Even so, Linnaeus did not always distinguish between mythological versus real crea-
tures and incredible versus credible hypotheses. His understanding of ‘cause and effect’ was circumscribed
by prevailing Lutheran theology and Cartesian mechanistic philosophy. Linnaeus was as much a working
physician, agriculturalist and land surveyor as he was a taxonomist. Contemporary economic biology and
biotechnology are anticipated in his animal and plant breeding and pearl culturing experiments. In the great
body of Linnaeus’ letters, manuscripts and books are discernable foundations for many other later disciplines.
These include: anthropology, biogeography, bioinformatics, biomechanics, biological control, conservation,
ecology, epidemiology, Darwinian evolution, ethnography, medical diagnostics, microbiology, palaeontology,
pharmacology and phylogenetic systematics.

KEYWORDS: biodiversity, evolution, Linnaeus, systematic, taxonomy

zoological classification (Jeffrey, 1973). Today there are

INTRODUCTION
The 18th century Swedish naturalist and physician
Carolus or Carl Linnaeus (1707–1778) was the main insti-
gator of the framework for biological taxonomy still in gen-
eral use today. This is the binary (binomial or ‘two word’)
Latin or Latinised system for scientifically recognising,
describing, naming and categorising genera and species
of plants and animals. Linnaeus in the first edition of Spe-
cies Plantarum (1 May 1753) attempted to document all of
the plant species then known. This is the accepted start-
ing point for validly published names in botany (Jeffrey,
1973). Through successive editions of Systema Naturae
(1735 onwards) Linnaeus consolidated and popularised his
method of nomenclature for both plant and animal species;
and the 10th edition (1 January 1758) is acknowledged as
the formal starting point for names available in modern
about twenty-five societies worldwide dedicated to hon-
our Linnaeus’ life and work. Among these, the Linnean
Society of London (www.linnean.org) contains his life-
time awards, some portraits, personal library, more than
3,000 letters and manuscripts and the bulk of his original
botanical and zoological specimen collections. These are
now being electronically catalogued, with web-accessible
digitised images. This is part of the Computerised Access
to the Records of the Linnean Society or CARLS Project
(Gardiner & Morris, 2007; http://www.linnean.org/index
.php?id=370).
It can be deduced from Linnaeus’ extensive collec-
tions of specimens, associated documentation, numerous
publications and voluminous personal correspondence
(Heller, 2007) that he was not only an inspired theoretician
but also a very practical man concerned to catalogue the

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TAXON 58 (1) • February 2009: 18–31
immense variety of life and express taxonomic relation-
ships in a logical, consistent, structured and stable way
(Hagberg, 1944; Blunt, 1971; Broberg, 2006; Harnesk,
2007; Schmitz & al., 2007; Hagelin, 2007). Fully adopted
everywhere by about 1778, fundamentals of the Linnaean
system of binomial nomenclature (including a nested hier-
archy of ranks to include class, order, genus, species and
variety) persist until now but with additional categories
(Stuessy, 2006). Nevertheless, there is a substantial and
growing academic challenge regarding the theoretical ba-
sis of the system and its future utility (Alves & al., 2007;
Mishler, 2009).
In a modern context, Linnaeus can fairly be regarded
as the originator of systematic and ecological studies in
biodiversity, having given Latinised names to some 7,700
species of plants and 4,400 species of animals (Dixon &
Brishammar, 2007). However, he greatly underestimated
the number of plant species in the world, confidently sug-
gesting less than 10,000 (Linnaeus, 1753) whereas modern
authorities propose that the total number of species of
vascular plants alone is near 420,000 (Govaerts, 2001;
Bramwell, 2002; Crane & Pleasants, 2006). Indeed, Lin-
naeus could hardly have imagined that the number of all
species of living organisms to be scientifically recognised
would eventually reach above 1.75 million, with perhaps
another 14 million yet to be described, according to the
Global Biodiversity Assessment (Watson & al., 1995 and
subsequent revisions). Nonetheless, the unifying and en-
during paradigm in natural philosophy forged by Lin-
naeus helped lay foundations for several major biological
disciplines developed in addition to taxonomy through
the 19th to 21st centuries. Overall, Linnaeus was great
communicator of practical and scientific knowledge. This
contrasts with earlier cultural and academic traditions
of tightly controlling the source, distribution and use of
knowledge and the language of communication. He often
popularised and radicalised his subject matter to gain at-
tention. For example, his sexual system for classifying
plants (Schmitz & al., 2007; Snoeijs, 2007) was conveyed
with a bawdy humour not acceptable to all of his peers.
He was also steadfast in placing humans with primates
in his classifications, so creating a great, productive and
continuing controversy (see below). In the Rudbeckian
tradition, public dissections of humans and other dem-
onstrations were organised in the University of Uppsala
(Lindroth, 1976).
He was particularly enthusiastic about his students
who often visited his private house and he gave inspir-
ing public presentations and private lectures or collegia
for a fee. He was good at obtaining funding for projects
and established an international research school of his
student ‘apostles’, demonstrating his personal magnet-
ism and entrepreneurial skills. He promoted the idea of
expeditionary research on his own account and through 17
Reid • Linnaeus’ life and relationship to modern biology
of the apostles. They surveyed nature at home and abroad
for both theoretical and practical or economic reasons
(Nordenstam & Lindell, 2007). In modern terms Linnaeus
can be regarded as a strong advocate for the public under-
standing of science, education and technology (PUSET).
The powerful Linnaean paradigm in taxonomy sprang
from his original and perceptive approach to understand-
ing and explaining the natural world. It also stemmed
from traditional beliefs and the environmental, economic,
demographic, social, cultural, political, theological, and
philosophical cum scientific influences of this and earlier
periods.
ENVIRONMENT AND ECONOMY
Born in 1707, Linnaeus grew up in the remote country
villages of Råshult and Växjö in the Swedish region of
Småland (Borglid, 2006; Broberg, 2006). Living condi-
tions were often demanding and harsh and there was direct
dependence on animals, plants and natural materials or
substances for food, clothing, shelter, goods and other
purposes. This direct dependence on nature is not seen in
21st century western industrialised societies. Only a low
percentage of the population now work close to the land
and utilise substantial amounts of wild-derived food and
other resources. This makes the survival imperatives of
this earlier generation difficult to fully appreciate. Any
failure to successfully engage with nature could have dire,
sometimes fatal consequences. Hence the recognition,
naming and classification of different kinds of animals,
plants and rocks — through oral traditions and informal
folk taxonomies — had pragmatic more than abstract di-
mensions. Knowing what was an edible wild plant or berry
and what was poisonous to a human or an animal (and any
seasonal changes) would have been essential, everyday
knowledge and vital to pass on to the next generation.
Linnaeus, with his acute powers of observation, or-
ganised mind and retentive memory must have been at a
particular advantage in this regard, even as a child. From
his youth onwards he was certainly fascinated by nature
in the broadest sense from stones to stars; and with the
many possible useful applications of this knowledge and
the great economic potential (Koerner, 1999). By stark
contrast, few students living today in westernised socie-
ties can accurately identify many species of wild animals
and plants or their products. This is a reflection of soci-
etal changes in perceived practical and economic needs.
There is an ill-conceived shift in emphasis and priorities
in modern biological science teaching, despite the mas-
sive challenges remaining (indicated above and below) in
documenting global biodiversity.
Key features of the physical environment in Lin-
naeus’ time were short summers, severe winters, frequent
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Reid • Linnaeus’ life and relationship to modern biology
agricultural fires, crop failures, famines, poor diet, inad-
equate sanitation, widespread epidemic disease and dental
problems. Short summers meant that there was only a
comparatively brief period (4–5 months) when life was
at its most obvious and exuberant; and when animal and
plants and their products needed to be quickly harvested,
processed and stored. This was in order to tide villagers
over a winter featuring poorly heated and inadequately
insulated houses, deep snow drifts and temperatures
probably commonly as low as minus 30°C. By contrast,
relatively mild winters have been experienced in recent
decades in northern Europe and elsewhere. This has been
scientifically attributed to human-induced climate change,
linked with actual and potentially deleterious impacts
on biodiversity. In this and other contexts the need for
taxonomic knowledge has never been greater (Dixon &
Brishammar, 2007).
Ironically, it was Linnaeus who in about 1743 devised
the reversed form of centigrade temperature scale devel-
oped in Uppsala University by his scientific colleague and
supporter Anders Celsius (1701–1744) (Lindroth, 1976).
Linnaeus recognised the value of the thermometer in hot-
house cultivation and used it on the 16 December 1745 in
the Orangery at Uppsala to produce the first ever tempera-
ture records. Subsequently, Linnaeus’ apostles recorded
temperature during natural history expeditions overseas.
Thermometers with the Linnaean scale are now in general
use and serve as a fundamental measuring device in the
physical, geographical, biological and medical sciences.
Sexually transmitted conditions, diseases of insanita-
tion and other potentially lethal epidemics were rife in 18th
century northern Europe (Singer & Underwood, 1962).
The ‘ague’, now known as malaria, was common — inci-
dentally becoming rare in the 19th and 20th centuries but
now returning. The spread is associated with a resurgence
of mosquito vectors and climate change. In the 18th cen-
tury the ague was of unknown cause and with no proven
cure: a subject upon which Linnaeus later conducted medi-
cal research, developed theories and experimented with
remedies (see below).
Frequent serious epidemics meant that individuals
needed to be physiologically robust to survive and there
was a high mortality, especially among infants. Linnaeus
himself was deeply affected by the neonatal loss of his
second daughter, born in 1744 and infant son Johannes,
born in 1754 (and who died in 1757). Untreated dental
decay was extremely common in the 18th century (Singer
& Underwood, 1962) and recent studies in developing
countries confirm that such decay is often not trivial and
can lead to fatalities, especially in the young. Portraits
of people throughout the 18th century are invariably un-
smiling, probably concealing dental disfigurement. Art
historians place a 1787 work by Elisabeth Vigeé-Lebrun
as the very first smiling portrait. Despite his genial nature,
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TAXON 58 (1) • February 2009: 18–31
Linnaeus was never depicted as smiling in any of several
portraits. He was plagued with dental problems and by the
age of sixty had no teeth (Blunt, 1971: 167). In his corre-
spondence he often grumbled over acute toothache — and
this unpleasant condition must have affected his disposi-
tion and capabilities throughout his working life. Unfor-
tunately, it was not until the close of the Linnaean period
that modern scientific dentistry was established by John
Hunter in The Natural History of the Human Teeth (1771)
and in A Practical Treatise on the Diseases of the Teeth
(1778). Here, knowledge gained through the compound
microscope was used to advantage. Recent biomedical
research (Lee & al., 2006; Hingorani & D’Aiuto, 2008) on
severe, chronic and cumulative periodontal disease (of the
kind very likely suffered by Linnaeus) indicates a strong
correlation with elevated systemic C-reactive protein. This
raised protein level is a response to the presence of vari-
ous periodontal bacterial pathogens whose toxic products
enter the bloodstream.
Excessive C-reactive proteins are, in turn, strongly as-
sociated with cardiovascular disease and cerebro-vascular
accidents (stroke), especially stroke history in the elderly.
In this connection it may well be significant that in old
age Linnaeus had angina in 1773, a severe semi-paralysing
stroke in May 1775, another in the winter of 1776/77 and
one finally on the 30 December 1777. This was shortly
followed by death on the 10 January 1778, aged 71 (with
his ashes buried under a flat gravestone inside the Cathe-
dral of Uppsala).
A central aspect of the wider demographic and social
environment that the young and maturing Linnaeus had to
operate and develop in was low human population levels.
In Råshult and Växjö the populations would have num-
bered in the hundreds at most and the entire population
of Sweden would only be about 1.5 million, in contrast to
today’s census figure of more than 9 million. Low popula-
tion numbers resulted not only from general plagues and
infant mortality (see above) but also because of fatalities
among soldiers in successive European wars including the
Thirty Years War (1618–1648) and its aftermath involving
various conflicts with the Danes, Russians, Saxons and
Poles. This led, in turn, to a shortage at home of young
males for physical and other tasks. The mean life expect-
ancy, from birth, of those born between 1755–1776 was
33.2 years for males and 35.7 for females—by comparison
to 73 or more years for both sexes today (Medawar &
Medawar, 1983: 68; Miles, 2006: 76). Conversely, this cre-
ated openings and a demand for employment, especially
for the better educated individuals. Nonetheless, educa-
tional and employment opportunities were limited and
Linnaeus was, therefore, fortunate to have been born into
a family of Lutheran parsons where formal education and
training for a career was valued and supported. Indeed,
the clergy in Sweden, historically had a privileged access
TAXON 58 (1) • February 2009: 18–31
to education. Sons of clergy made up as much as 42% of
the student body in Uppsala in the mid-17th century and
as many as 65% of graduates entered the service of the
Church (Lindroth, 1976).
Linnaeus first went to a grammar school in Växjö
where his science teacher, the state physician Johan Roth-
man, encouraged the study of botany. Rothman proposed
to Linnaeus’ parents that he should train for a career in
medicine rather than carry on with the family tradition
of studying theology and becoming a parson (Broberg,
2006). From Växjö Linnaeus moved to the University
of Lund in the province of Skåne. In 1728 he went on
to study in the University of Uppsala further north near
Stockholm in the province of Uppland. Eventually, in a
hotly-contested competition of 1741, he secured the posi-
tion of Professor of Medicine and Botany (Blunt, 1971;
Lindroth, 1976: 96, 97; Broberg, 2006: 44).
SOCIETY AND CULTURE
For the future and lasting benefit of taxonomy it is
surely no coincidence that the patrilineal (‘son of’) system
of naming ended around the time of Linnaeus’ birth in
1707. Up to this point, the family name had been lost with
each successive generation. Had things been otherwise,
Linnaeus would simply have adopted the first name of
his father Nils Ingemarsson and been called Carl Nilsson;
with his own sons, in turn, being given the family name
Carlsson. The patrilineal system sufficed in the 16th and
17th centuries because everyone would have known each
other in villages. There were small populations and vil-
lage inhabitants were not geographically mobile. A bet-
ter system of individual identification became necessary
with population increase and the more complex social,
legal, educational, employment and travelling arrange-
ments resulting from expansion of villages and develop-
ment of towns and cities in the 18th century. Interest-
ingly, Linnaeus’ name derives from the Latinisation of
‘Linn’— the name in Småland dialect for the Linden or
Small-leaved Lime tree (Tilia cordata Mill.). This is a
widespread northern European species which would have
been seen everyday by his parents and grandparents and
which is pharmacologically and economically useful even
today. Crucially, this new system for naming humans was
often inspired by nature and allowed fresh perspectives
in tracing human genealogies and expressing ancestor-
descendant relationships, eventually including plant and
animal breeding records and taxonomic classifications
(see below).
Linnaeus had modest social origins and thus faced
great barriers to educational and career progression from
a well-defined social hierarchy, tightly controlled military
governorships of regions, still higher royal prerogatives
Reid • Linnaeus’ life and relationship to modern biology
and a complete dependence on patronage at every stage.
These potential impediments were all skilfully negotiated
by Linnaeus, evidently a man of great charm and cha-
risma. He achieved advancement through a socially astute
marriage to Sara Lisa Morea and also through the patron-
age of King Adolf Fredrik I and Queen Lovisa Ulrika of
Sweden. They made him Court Physician and Curator of
the Royal Natural History Specimen Collections (Reid,
2007). As the ultimate reward, he was ennobled by them
in 1757, gaining the Teutonic title Carl von Linné.
As a young man Linnaeus traveled widely in Sweden
and later in Western Europe for the purposes of botanical,
zoological, geographical and geological study and for the
economic advancement of himself and others (Koerner,
1999). A lack of maps, the short summer season, difficult
terrain (often without paths) and the frequent presence
of robbers meant that Linnaeus’ early travels in Sweden
were not always easy. Nonetheless, he published thorough
accounts of his Lapland then Öland and Gotland journeys
conducted at the ages of 25 and 34, respectively (Black &
Lee, 1979). These incorporate ethnographic descriptions,
well-reflected in the famous painting by Martin Hoffman
(1737) of a young Linnaeus dressed in traditional (Saami
female!) Lappish regalia.
Swedish land surveying and cartography had already
been well established to a high standard in the mid-17th
century by Andreas Bureus (1571–1646) as shown in the
copper engraving Svecia, Dania et Norvegia, Regna Euro-
pae Septentrionalia (published ca. 1640). However, in the
late 17th and early 18th centuries access to maps in Sweden
and elsewhere in Europe was strictly controlled by regional
governors and military planners. Maps were very costly to
produce and print and represented a privileged military,
economic and political knowledge. This was a power to
be used to advantage, including for national propaganda,
legitimising the state and for minimising dissent. Hence,
in Linnaeus’ youth, maps were not generally accessible,
relied on received wisdom and ancient, outdated informa-
tion. Maps had large gaps, a fair level of inaccuracy and
some mythology—being rhetorical exercises in Latin more
than empirical studies (Legnér, 2001).
During Linnaeus’ time, and partly as a result of his
endeavours, there came an ‘Age of Liberty’ or ‘Age of
Freedom’ in Sweden (ca. 1720–70) — a local aspect of a
general European and North American philosophical and
scientific ‘Enlightenment’, which sprang in turn from the
17th century ‘Age of Reason’ (Taton, 1964; Russell, 1992).
In the Age of Reason “instruction and learning were still
dominated by Lutheran theology and Roman eloquence”
(Lindroth, 1976: 92, 93). The new Age of Freedom pro-
duced a general reaction against the use of dead languages
in communication and against the teaching of an abstract
metaphysics concerning principles of reality (Dixon &
Brishammar, 2007: 150). Supported by Linnaeus and
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Reid • Linnaeus’ life and relationship to modern biology
others, including Abraham Hülphers, geographical de-
scriptions published post 1741 were of far better quality,
more numerous and more widely available. Significantly,
Swedish rather than Latin became the language of expres-
sion (Legnér, 2001: 2).
Reflecting society and culture, there is something
of a military element in the plans, charts, sketches and
maps of Linnaeus’ early studies, exemplified in his note-
book or Örtaboken of 1725. His published taxonomies
perhaps mirror the military hierarchy and the divisions
and levels at which soldiers were drawn into service from
village and district to province and kingdom (Broberg,
2006: 18). Indeed, he once jokingly classified his botani-
cal colleagues in a ranked military system with himself
as the commanding general (Nordenskiöld, 1920: 212).
This militaristic mindset coupled with theological and
scientific influences has, it seems, been transmitted for-
ward into medical diagnostics and modern hierarchical
expressions of phenetic, morphological, geographical and
phylogenetic relationships.
Whatever its origins, Linnaean geography can be seen
as an essential precursor to the scientific study of what
individual organisms, populations, species and higher-
level taxa occur where and why. This is now expressed as
distributional maps of plants and animals in ecology and
historical biogeography, including vicariance analyses;
and as epidemiology in modern human and veterinary
medicine. The familiar and reassuring ‘May Tree’ and
other diagrams of Linnaeus, with more-or-less dichoto-
mous (binary) branching, precede modern expressions of
taxonomic or phyletic relationship such as phenograms
and cladograms — whether morphological, palaeo-geolog-
ical or palaeo-geographical. In this aspect Linnaeus was
probably influenced by the ancient Norse or Germanic
Yggdrasil, a mystical World Tree of Life or axis mundi. A
similar concept appears in antiquity in many other cultures
and in the Genesis origin myth in the Christian bible. This
concept is often associated with the idea of a scala naturae
or ‘Great Chain of Being’ (Schmitz & al., 2007: 29) and
other pagan, theological or quasi-genealogical concepts
of descent. Geographical data, when combined with geo-
logical, taxonomic, genealogical and genetic data from
the mid-19th century onwards, eventually produced the
profound evolutionary framework of space, time and form
that we recognise today (see below). For Linnaeus’ ad-
mirer Erasmus Darwin (1794) the Great Chain of Being
somehow expressed in deistic terms the morphological
transformations or ‘evolutionary’ series that he observed
across taxa. For his grandson Charles Darwin (1859) the
Tree of Life represented an agnostic means by which the
origin and evolution of species and higher-level taxa could
be generalised (see below). Mishler (2009) evidently sees
the Tree as a derived cladistic concept, somehow dissoci-
ated from its original roots.
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TAXON 58 (1) • February 2009: 18–31
THEOLOGY AND TRADITIONAL
BELIEFS
Linnaeus lived in a staunchly Lutheran theocracy
where the protestant ethic was all pervasive, including
an imperative to be thrifty, work hard and with an em-
phasis on educational attainment. This, together with the
moderating influence of the Age of Reason, was very
important for the development of his critical thinking.
Within this, came the fostering of a spirit of quiet con-
templation and individual free enquiry — without the Ro-
man Catholic necessity of using priests as intermediaries
to gain spiritual and other insights into the workings of
the world. In pursuing Lutheran knowledge, there was
also an emphasis on the written word and book learning
over ‘graven images’ and an impetus to determine and
celebrate God’s Plan and communicate this knowledge
far and wide.
Linnaeus was brought up in a family home and wider
society where strict religious observance was the norm
and where, even in a university setting, theologians had
primacy (Lindroth, 1976). From this, there were definite
social, cultural and career limitations on what could be
learned and said in the name of science (a situation per-
sisting to a greater-or-lesser extent today in many human
societies, including westernised ones). Despite an outward
appearance of piety, Linnaeus was in many respects a
rationalist unafraid to challenge accepted norms. This
disposition could have cost him his academic position at
Uppsala University. Certainly, in the wake of the Enlight-
enment and with the growth of science and utilitarianism,
“Linnaeus’s orthodoxy was doubted by his theological
colleagues” (Lindroth, 1976: 123) and he was accused of
impiety by the Lutheran Archbishop of Uppsala. In this
sense he was a religious non-conformist, or ‘free thinker’.
Professional development involved travel to both the Neth-
erlands and England (1735–1738). These countries shared
the same sort of radical protestant ethic and likely rein-
forced it in Linnaeus.
Linnaeus’ scientific heroes and immediate predeces-
sors were often devoutly protestant, yet spiritually and
intellectually deviant — including Robert Boyle and John
Ray, two 17th century British polymaths with botanical
interests who espoused a particular Natural Theology.
From this, Linnaeus became increasingly drawn to explor-
ing nature’s potential benefits to man. It was clear from
Boyle, Ray and others that God had created the earth and
its bounty for man to explore, explain and utilise (Koerner,
1999). Linnaeus also greatly admired Sir Isaac Newton,
another deeply religious individual, biblical scholar and
natural philosopher. Newton incessantly challenged basic
assumptions about the nature of the universe and (de-
spite protestations to the contrary) developed the experi-
mental method in science, with the system of competing
TAXON 58 (1) • February 2009: 18–31
hypotheses. He gave full expression to this in Principia
Mathematica (1687). Indeed Linnaeus had aspirations to
be seen as the intellectual equivalent of Newton, with
the Systema somehow securing for botany and zoology
the physical and chemical profundity and success of the
Principia — a work which Linnaeus admired but evidently
had difficulty in understanding (Broberg, 2006: 35). Of
course Linnaeus’ work, while cataloguing excellent facts
gained through direct observation and making broad and
valuable biological generalisations, lacks the extraordi-
nary breadth, depth and explanatory power of Newton’s
physical laws.
Despite a certain religious unorthodoxy, numerous
deistic quotes are ascribed to Linnaeus the most famous
one being: “Deus creavit, Linnaeus disposuit ” — God
creates, Linnaeus disposes (for this and other examples
see Landström & Modin, 2006). This may or may not,
given the prevailing social and religious constraints, dem-
onstrate his true love of God and belief in the Creation.
Wolff ianism, a modified creationist doctrine promulgated
by the German philosopher Christian von Wolff, proposed
a natural theology for the Enlightenment centred on
“physicotheology” which held that an omniscient Creator
could be logically deduced from the order and beauty of a
Creation governed by physical and natural laws (Schmitz
& al., 2007: 28). In Uppsala this idea was “elevated by
Linnaeus into a touching religious devotion” (Lindroth,
1976: 128). Elements of Wolffianism persist today in the
widespread anti-scientific and anti-evolutionary Chris-
tian fundamentalist argument for ‘intelligent design’, now
undergoing a modern creationist revival in westernised
society, particularly the U.S.A.
In 1736 Linnaeus met with Sir Hans Sloane, a con-
temporary of Ray and the founder of the British Museum.
Sloane’s inspiration affected many scholars and also led
to the establishment of the British Museum (Natural His-
tory), now the Natural History Museum, London. The
libraries of both institutions now contain many Linnaean
publications (British Museum, 1998). A close identifica-
tion with the high achievements, thinking, prestige and
authority of his philosophical and scientific forebears evi-
dently meant a great deal to Linnaeus. Such historical and
contemporary influences must have played a large part in
shaping and regulating his views. The taxonomic princi-
ple of authority and precedence that he developed — even
now strongly embedded in modern systematic biology, but
perhaps beginning to wane — probably has theological and
military as well as scientific roots. The biblical creation
myth of the original and representative humans ‘Adam and
Eve’ may in Linnaean taxonomy possibly be reflected in
the development of a ‘typological’ thinking where indi-
vidual specimens are held to represent fixed and unchang-
ing species and genera. This has historically and perhaps
erroneously been linked to Aristotelian essentialism
Reid • Linnaeus’ life and relationship to modern biology
and its supposed intellectually stultifying effects on the
progress of taxonomy (Nordenskiöld, 1920: 207; Hull,
1965; Schmitz & al., 2007: 28; Winsor, 2006a, b).
Folklore and superstition sometimes became con-
fused with theology and science in Linnaeus’ thinking
and he did not always distinguish between real and myth-
ological creatures. This is illustrated in the classically
influenced ‘Paradoxa’ sections of the 1st to 8th editions
of the Systema. From Nordic sagas he probably believed
in trolls, was fascinated by contemporary women folk-
healers claiming to be witches and would include hydra,
mermaids and other fabulous taxa in his classifications, at
least as possible species. Broberg (2006: 12) finds that:
The early medical studies reflected by Linnaeus’ notes
from this period present a remarkable blend of traditional
belief or superstition and modern mechanistic medicine.
In Linnaeus’ early life and work there is a definite
contrast between the rational approach to understand-
ing problems and a reliance on irrational, unscientific
folk traditions and myths. Newtonian principles of cause
and effect, with logical sequences of reasoning, were not
always followed. Nonetheless, in Uppsala he soon sub-
scribed to the ‘mechanistic philosophy’ of René Descartes
(1596–1650); later debated by Jean-Jaques Rosseau (1754)
and others. The basic proposition is that man is an animal
with anatomical and physiological functions analogous to
a machine. This far-reaching scientific model (still with
us today, translated as biomechanics), was adopted by
Linnaeus as: “Homo est animal ” and “Homo machina
est ” (Broberg, 2006: 13). Indeed, for his doctoral thesis
on ague — successfully defended in Harderwijk University
in the Netherlands in 1735 — he followed biomechanical
arguments. Ingested clay was postulated to be a causal
agent for the ague, somehow clogging up the human ma-
chine to create the symptoms. Contrary to this, he treated
his sister in Stenbrohult who was suffering from ague
with a strange, unproven folk remedy by wrapping her in
raw, freshly slaughtered sheepskin. In any event, Johan
Gottschalk Wallerius considered Linnaeus’ doctoral thesis
on the ague to be mediocre. He challenged him dramati-
cally for the Professorship of Medicine in 1741 but lost
(Lindroth, 1976).
Even in straightforward matters of natural history,
Linnaeus sometimes had difficulty in establishing credi-
ble versus incredible hypotheses. For example, when faced
with the puzzling question of where the small passerine
Barn swallows (Hirundo rustica L. 1758) go in winter he
imagined that they dived to the muddy bottom of ponds to
over-winter. This contradicts what he may, could or should
have known by that time in terms of basic physiology and
the seasonal migration patterns of birds. In this instance
Broberg (2006: 12) considers that:
23
Reid • Linnaeus’ life and relationship to modern biology
… one is entitled to demand better things of an academic
naturalist … Mistakes like this may seem embarrassing,
but they are more appropriately to be regarded as tensions
in a culture of many different strata.
While not a shining moment in science, there is to
Linnaeus’ credit circumstantial and observational evi-
dence that swallows fly low over ponds and become ab-
sent. The entire hypothesis of hibernation underwater is
at least potentially testable through pond draining and
digging and experiments on swallows.
APPLIED KNOWLEDGE
The Age of Freedom in society and culture coincided
with a Swedish government policy where the economy in
terms of industry and trade had the highest priority. Un-
der this regime the applied physical and natural sciences
flourished at the University of Uppsala (Russell, 1992: 88)
and, indeed, continue to prosper today. Linnaeus was as
much a physician, agriculturalist, land surveyor and gen-
erally practical man as he was a taxonomist. Many areas of
modern medicine, pharmacology, applied biology and bio-
nomics are indicated in his work. We know that Linnaeus
and his colleagues enjoyed imbibing alcohol socially from
correspondence to that effect. Interestingly, the pioneering
large-scale, modern biotechnology industry Amersham
Pharmacia Biotec — based close to Linnaeus’ house in
Hammarby near Uppsala and working in collaboration
with the University — has brewing research as one area of
economic production. Today, biotechnologists are, among
many other things, working on methods to genetically
engineer useful synthetic bacteria (Mycoplasma) as ‘new
species’. Depending on the modification, these might be
capable of producing green fuels to replace coal and oil,
digesting toxic waste or absorbing greenhouse gases.
Linnaeus began his life as a working physician, where
he developed a good reputation in the traditional study
and use of natural, often plant-derived drugs and he con-
tributed to various pharmacopoeias of ancient Greek,
Arabic and other derivation. These contained supersti-
tious and sometimes repugnant folk remedies as well as
efficacious, if crude, substances including mercury prepa-
rations possibly of Indian origin (Singer & Underwood,
1962: 673). Linnaeus successfully used the latter in the
treatment of venereal disease, so building up his medical
reputation (Blunt, 1971: 131). However, he may not have
used the compound microscope habitually and intensively
(Brian Ford, pers. comm.). Had there been a germ theory
of disease in the 18th century, this would have been an
excellent tool in terms of his doctoral study of the ague
(or malaria, see above). Understandably, in the absence
of an alternative microbiological framework, Linnaeus
24
TAXON 58 (1) • February 2009: 18–31
adopted a gross biomechanical approach to understanding
the disease. This was a false trail because Plasmodium is
the microscopic parasite now understood to cause malaria.
In principle, it would have been visible under compound
microscopes available to Linnaeus. These were techni-
cally well advanced by the 18th century (Hogg, 1882).
Linnaeus’ Lapland journey (1732) as a young man,
seems to have initiated the ecologist and economic biolo-
gist in him as well as engaging him in geography, min-
eralogy and the ethnology of the Saami Lapps. During
this journey, Linnaeus’ attention was drawn to natural
freshwater pearl fisheries where he noted the waste in de-
stroying 2,000 mussels to get one pearl. Adopting Chinese
techniques, he later succeeded in using chalk granules
to seed pearls in the mantle of bivalve molluscs such as
Angel Wings Cyrtopleura costata (L. 1758) and harvested
them successfully in the 1740s. These cultivated pearls are
externally indistinguishable from spherical wild pearls.
Original specimens exist in the collections of the Lin-
nean Society of London. Some of these have, since 2001,
been publicly exhibited on a world tour of the “Pearls”
exhibition, organised and administered by the American
Museum of Natural History, New York. This Linnaean
biotechnology helped give rise to an immensely valuable
modern global industry.
Influenced by Ray, Boyle and others (see above) he
became wedded to the concept of a ‘divine economy of na-
ture’ where everything has a use and is ultimately recycled,
e.g., domestic waste is eaten by animals, then animals are
eaten by humans (Koerner, 1999). Nature is also seen as
‘self-regulating’. In modern terms this can be expressed
as a feedback-governed equilibrium, including for the car-
bon cycle and the much publicised ‘carbon footprint’ of
humans. Such concepts are central to modern biotechnol-
ogy, applied ecology and sustainability and the notion of
‘ecosystem services’ in relation to human development
and poverty alleviation. Linnaeus expressed a concern for
practical nature conservation in his correspondence on the
negative impact of slash and burn agriculture on Finnish
lichen meadows and birch woods. He also noted goats over-
grazing on Swedish heather moors and sand dunes and
the general destruction by logging of northern European
forests. He considered the latter to be a temporary and
reversible blunder (Koerner, 1999: 85). By contrast, his
apostle Daniel Rolander (1723–1793) suggested in a letter
to Linnaeus (11 July 1755, Linnean Society correspondence
reference L1932, translated from the original Swedish) that
the South American rainforests in Surinam were so vast
that they could never be depleted and that, “humans, fire
and other things” have no appreciable impact:
If they [the insects] do not do it, then I am sure that this
huge Hothouse will stand un-mutilated as long as the
earth itself. [Encouragingly, we know today that 96.2% of
TAXON 58 (1) • February 2009: 18–31
primary forest in Surinam still exists (http://rainforests.
mongabay.com/deforestation/2000/Suriname.htm)]
Linnaeus’ agricultural economics concerned coop-
erating with rather than battling nature (Koerner, 1999:
83) — again, a guiding principle in modern applied ecol-
ogy, pest control and biotechnology. Interestingly, but
separately from notions of ague (see above), Linnaeus
proposed the idea of exterminating pest insects using
other insects. This is a forerunner of modern methods in
biological control, including for crop pests and malarial
mosquitoes. Linnaeus was particularly concerned over
crop failures and the economic and human consequences
and (in modern terms) the technological underdevelop-
ment that allowed for failures. From this he developed
an acclimatisation project to ‘teach’ tea, saffron and rice
to grow in the Arctic Tundra and some have found this
attempt to be bizarre. Instead, these endeavours can be
interpreted as a Linnaean intuition well ahead of its time,
anticipating the modern genetic engineering of crops to
cope with extreme environments. Nonetheless, his pro-
pensity to introduce exotic animals and plants into Sweden
to determine their practical use also presages present-day
European and global conservation problems associated
with the deleterious impacts of invasive ‘alien’ species.
Linnaeus was interested in the domestication and
breeding of animals, which can be regarded as a tradi-
tional biotechnology. He attempted, through experiments,
to domesticate yak, guinea pig and European elk as Swed-
ish farm animals. However, the science of genetics was
yet to be developed and so he did not have a good under-
standing of mammalian inheritance. This is evidenced in
his odd ideas on male genital melanism, which he argued
would appear in offspring resulting from crossing a hu-
man male negro with a Caucasian female (Broberg, 2006:
12). However, Linnaeus did realise that artificial selection
and hybridisation of livestock could bring about a change
or variation of appearance in offspring. This is significant
in terms of his later, modified views on the fixity of spe-
cies. Recognition of variability and ‘plasticity’ in species
would be essential to the later development of the Darwin-
ian theory of evolution by natural selection (see below).
LINNAEAN SYSTEMATICS
Aristotelian ‘essentialism’ — a view that any particular
entity in substance or form has a defining set of essential,
fixed and all-inclusive characteristics — may have domi-
nated pre-Darwinian systematic thinking and supposedly
had intellectually stultifying effects (Nordenskiöld, 1920;
Hull, 1965). Shared derived taxonomic characters (ho-
mologies, synapomorphies or synapotypies) operating at a
set level of generality in diagnosis might, in contemporary
Reid • Linnaeus’ life and relationship to modern biology
terms, actually constitute the elusive ‘essence’ that Aris-
totle was pursuing. Be that as it may, Winsor (2006a, b)
advances cogent arguments to demonstrate that Linnaeus
was not an ‘essentialist’. From Lindroth’s (1976) detailed
history we know that Aristotelian logic and scholasticism
was the dominant philosophy at the University of Uppsala
by the mid 17th century and that Aristotle (the founding
father of systematic zoology) was “regarded by the Lu-
theran dogmatists as an indispensable ally” from his strict
order and clarity of thought and the system of definitions
provided. Indeed, it has been strongly argued that Lu-
theran-backed Aristotelianism was the major inhibitor of
all Swedish science during the 16th and 17th centuries. At
this time, anti-Aristotelian writing might well be viewed
as “a form of intellectual insurrection … tantamount to
an assault on Lutheranism” (Russell, 1992: 87).
By the time Linnaeus arrived in Uppsala, Aristote-
lian philosophy had largely given way to the Cartesian
mechanical model for interpreting the world (see above).
There was evidently a “last autumnal flowering” of Ar-
istotelianism in Uppsala (Lindroth, 1976: 72) but there is
no reason to assume that Linnaeus fell unduly under its
influence, even if he acknowledged its original benefits
and utilised some of the reasoning. The major proponents
of the new Cartesian mechanics in Uppsala were the Pro-
fessors of Medicine, Hoffvenius and Rudbeck who had
been won over to this doctrine during earlier studies at
Leyden in the Netherlands. Olof Rudbeck the Elder was
a distinguished anatomist and botanist who established
a botanical garden in Uppsala at Svärtbacken. It fell into
disuse and was eventually restored to its former gran-
deur by Linnaeus. Olof Rudbeck the Younger travelled
in Lapland and collected and drew plants and animals,
becoming the founder of zoology and anatomy in Uppsala,
and another mentor to inspire Linnaeus and encourage
him in mechanical philosophy and expeditionary work.
Expressing the Rudbeckian tradition in researching com-
parative anatomy, Linnaeus displayed an inspirational use
of morphological characters for describing taxa. This was
notably expressed in the artificial sexual system of clas-
sification that he developed for flowering plants (Schmitz
& al., 2007; Snoeijs, 2007).
In 18th century Sweden, according to Lindroth (1976:
100):
Scientific research had come to the universities to stay; the
promotion of natural sciences by performing experiments
and making observations was just as important as lectur-
ing and giving collegia. No one was more insistent on this
point than Linnaeus.
Certainly, in the main body of his work Linnaeus
clearly demonstrates the benefits of a new philosophy and
adopts a progressively more rigorous scientific approach.
25
Reid • Linnaeus’ life and relationship to modern biology
There is a careful gathering of information through travel
and exploration, hypotheses, empiricism, accurate ob-
servations, comparisons, dissections, measurements and
counts (e.g., of the number of floral parts). He adopted
contemporary methods in establishing facts, including a
hand lens and pocket microscope in the field and probably,
to a far lesser extent, a compound microscope in the labo-
ratory. Nonetheless, later editions of the Systema Naturae
notably incorporate microorganisms and therefore ‘set the
stage’ for modern microbiology. He also makes botanical
and zoological generalisations concerning species and
genera; and organises knowledge in the sort of systematic
and hierarchical way essential for taxonomic and other sci-
entific schemes, insightfully separating designation from
description. The creation of a logical, heuristic framework
within which to easily insert new taxonomic information
anticipates modern ‘bioinformatic’ computerised systems
for bulk data handling. Pledge (1966: 89) observes that:
“Linnaeus was the first man to be able to cope with the
flood of extra-European species.” Finally, there are the
widely disseminated published descriptions, diagnoses,
explanatory diagrams and literature reviews which make
his science historically well-informed, publicly accessible,
challengeable, testable and to an extent predictive.
Crucially for science and taxonomy there is an implicit
Linnaean requirement for material evidence to support
hypotheses. His specimens of animals and plants were
probably originally intended to simply be ‘name bearers’
for species (the stated purpose of types in the modern
International Code of Botanical Nomenclature, Article
7.2). Nonetheless, Linnaean specimens (which were not
then viewed as types in the contemporary sense of the
ICBN) are representative evidence for the recognition of
entire species and higher-level taxa. Specimens consti-
tute fixed reference points in any disputes on identity, or
over the morphological characters used in diagnoses and
differentiation. The Prefect’s House erected by Rudbeck
in one corner of the Uppsala Botanical Garden became
Linnaeus’s home for much of his life and there he kept his
herbarium and collections of shells, insects, fishes, fossils
and minerals. In 1768 he purchased a country residence
nearby in Hammarby and moved his museum to a small
outbuilding on the hill above. It is this collection which, in
large part, has ended up being preserved by the Linnean
Society of London (Gardiner & Morris, 2007).
Linnaeus’ techniques for specimen preservation were
limited and his process of drying, dissection and storage
(which relate to that used domestically for over-wintering
human and animal food) resulted in a subsequent deterio-
ration and loss of valuable biological evidence, especially
in fishes and molluscs. From simple considerations of
space and available technology, this method was also ar-
bitrary and highly selective. For example, only specimens
that could be dried were kept, and larger specimens could
26
TAXON 58 (1) • February 2009: 18–31
not be accommodated, nor large numbers of specimens
(Gardiner & Morris, 2007). Curiously, the useful meth-
ods of wet preservation of specimens in brine or alcohol
(known in Europe since at least 1662; Reid, 1994: 32, 33)
seems not to have been used extensively by Linnaeus, if
at all. This may well be because of the prohibitive cost of
preservative spirit and glass containers.
In today’s terms, the specimen sample sizes of Lin-
naeus are too small to properly characterize natural varia-
tion in populations, morphological or otherwise. To prove
useful, biological concepts in the diagnosis of species ulti-
mately need to express such variation through representa-
tive and statistically significant samples of populations.
More recent ideas of organic evolution, phylogeny and
genetics depend on an understanding of biological varia-
tion (see below). The Linnaean implication that the species
is the fundamental unit in diagnosis and that one or a small
number of specimens are adequate to diagnose species
has led to an effective, if procedurally elaborate, modern
system. However this system still places undue reliance
on one or only a few ‘type’ specimens as name-bearers,
vouchers or standard references for species. This is despite
the fact that such types may be biologically unrepresenta-
tive of wild populations and were often collected in the
18th and 19th centuries without good locality and other
supporting data. The typological approach became for-
mally enshrined in the 19th century following Alphonse
de Candolle’s 1867 attempt to formalise rules for botanical
names. This eventually led to the establishment of the
International Code of Botanical Nomenclature, 1930, and
the International Code of Zoological Nomenclature, 1895
(Melville, 1995; Pavord, 2005; Stuessy, 2006). A debate
continues over the nature, origin and utility of the concept
of a type and the ‘typological’ approach and whether or
not it is time to abandon it altogether (Winsor, 2006a, b;
Alves & al., 2007; Mishler, 2009).
Linnaeus collected specimens himself in Europe and
also received them from colleagues and students. These
specimens sometimes came from abroad, together with
descriptions and notes. Geographical and international as-
pects of scientific exploration were certainly well catered
for through Linnaeus’ seventeen most widely travelled
students, or ‘apostles’. Expeditionary research was, for
example, conducted in North America by Pehr Kalm, in
South America by Pehr Löfling and Daniel Rolander, in
China by Pehr Osbeck and in Arabia by Pehr Forsskål.
Also, Daniel Solander and Anders Sparrman sailed with
Captain James Cook around the world as his ship’s nat-
uralists (Lindroth, 1976; Gardiner & Morris, 2007). A
particular favourite of Linnaeus was Forsskål who was
born in 1732 in Helsingfors, Finland and educated both
in Uppsala University and in Göttingen University (in
present-day Germany). Interestingly, Forsskål was an out-
spoken critic of Wolffianism (see above) and also of Johan
TAXON 58 (1) • February 2009: 18–31
Gottschalk Wallerius, the principle advocate of this theo-
logical system in Uppsala University (Lindroth, 1976).
Wallerius, who lost academic preferment in competition
against Linnaeus, probably bore lingering resentments
against Linnaeus and, by proxy, his most heretical student.
At Linnaeus’ suggestion, Forsskål joined a Danish expedi-
tion to Arabia in 1761 and was the first person to describe
the unusual plant and animal life of the Red Sea. Sadly,
he died of malaria in the Yemen in 1763. Linnaeus was
somehow held responsible for sending the young man to
his death (Reid, 2007).
NATURAL RELATIONSHIPS AND
EVOLUTION
In developing the binary system, Linnaeus drew in-
spiration from many sources notably including his imme-
diate predecessors John Ray, 1628–1705, Joseph Pitton de
Tournefort, 1656–1708 and his contemporary at Uppsala
Pehr Artedi or Pectrus Arctaedius, 1705–1735, whose
Ichthyologia (1738) was published posthumously by Lin-
naeus (Nordenskiöld, 1920–1924; Dixon & Brishammar,
2007; Gardiner & Morris, 2007; Reid 2007).
Ray’s remarkably prescient concept of a species was
of an interbreeding population, thus introducing the (still
hotly debated) idea of biological process being used to
define species, as apposed to form and pattern alone.
However, this entity was deemed to be created and fixed
by God, albeit with evidence for transitional series of
taxa and other anomalies that were difficult to explain
(De Beer, 1970: 1; Pavord, 2005: 384). At first, Linnaeus
evidently accepted these arguments only in part, arguing
that both genera and species were objective realities, fixed
and immutable:
Species tot numeramus quot diversae formae in principio
sunt creatae [We enumerate as many species of diverse
appearance as were created in the beginning]. (Linnaeus
in his introduction to Philosophia Botanica, 1751: 99)
In the last (1768) edition of Systema Naturae Lin-
naeus gave a complete list of fossils in the order of their
succession, implicitly recognising them as representing
animals and plants that were no longer extant. Hence,
he laid the foundations of palaeontological classification
(Pledge, 1966: 90) and evolutionary stratigraphy. He also
eventually allowed that, while living genera remained
fixed, there might be change within or between species,
perhaps through hybridisation (Nordenskiöld, 1920: 214;
Pledge, 1966: 90; De Beer, 1970: 1). By contrast, the
French naturalist Buffon (1707–88) considered that only
individuals were objective realities, all other taxonomic
categories being human abstractions (Nordenskiöld,
Reid • Linnaeus’ life and relationship to modern biology
1920–1924: 224). This profound and contentious debate
is so far unresolved. The fact that Linnaeus (1753) used
the word relatus in for example the introduction to his
Species Plantarum and often espoused the desirability
of developing a ‘natural’ system of classification, does
not mean that he had a true genealogical or phylogenetic
sense of systematic relationships. Ideas of individuals,
species, genera, families and taxonomic hierarchies were
not necessarily organic, i.e., fully imbued with a sense of
life, reproduction, inheritance, descendants and ancestors.
Certainly, he applied the same classificatory scheme and
terms to both biological specimens and inorganic geologi-
cal samples. However, natural variation (with the prospect
of natural selection acting on this) is perhaps presaged in
18th century sketches (apparently Linnaean) of the diverse
beaks of tropical birds. These sketches are in the exhibited
collection of the Uppsala University Museum and they
look remarkably like those Charles Darwin made, much
later on the voyage of the Beagle (1834–36).
In any event, determining organic relationships is the
key to later concepts of Darwinian evolution, phylogeny,
and biogeography. New perspectives in expressing hu-
man family relationships probably allowed Linnaeus to
systematically recognise four human races (which even-
tually precipitated the modern biological, psychological,
sociological and political controversies on the issue of
‘race’). Linnaeus’ observations of non-human primates
were based on his own living menagerie including an
Orang-utan and on anatomical material. Crucially, this
led him to determine close affinities between humans,
great apes and monkeys. The idea proved to be exceed-
ingly controversial at the time and continues to be so:
The unbounded dominion which Linnaeus has assumed
in the animal kingdom must upon the whole be abhorrent
to many … He can hardly forbear to make man a monkey
or the monkey a man. (Albrecht von Haller in a review of
Linnaeus’ Fauna Suecica, 1746 in Blunt, 1971: 123)
It is not pleasing to me that I must place humans among
the primates, but man is intimately familiar with himself.
Let’s not quibble over words. It will be the same to me
whatever name is applied. But I desperately seek from you
and from the whole world a general difference between
men and simians from the principles of Natural History.
I certainly know of none. If only someone might tell me
one! If I called man a simian or vice versa I would bring
together all the theologians against me. Perhaps I ought
to, in accordance with the law of the discipline [of Natural
History]. (Letter from Linnaeus to Johan Georg Gmelin,
February 25, 1747, in The Linnaean Correspondence:
http://linnaeus.c18.net/mss_combine/UUB/L-GmelinJG/
L0783-a-150-02.jpg; http://linnaeus.c18.net/Letters/dis
play_txt.php?id_letter=L0783)
27
Reid • Linnaeus’ life and relationship to modern biology
I reject [Linnaeus’] first division, which he calls Primates
because my vanity will not suffer me to rank humankind
with apes, monkeys, and bats. (Thomas Pennant, 1771)
Recently many of our best naturalists have recurred to the
view first propounded by Linnaeus, so remarkable for his
sagacity, and have placed man … under the title of the Pri-
mates. (Charles Darwin in the Descent of Man, 1874: 230)
No anthropologist believes humans descended from
present-day species of monkeys or apes (Hurd, 1981).
Following Rudbeckian and Linnaean principles of
comparative and functional anatomy and only a year after
Linnaeus’ death, the German scholar Johann Blumenbach
(1779) moved to an approach where, on basis of upright
gait, he separated humans off from the rest of primates
(apes and monkeys). Some authorities today on shared
specialised (synapotypic) characters and cladistic argu-
ments would rather group chimpanzees and bonobos to-
gether with humans in the Linnaean genus Homo (Good-
man & al., 2002; Reid, 2002) — a conclusion still far too
radical for some.
In the same year as Darwin published his Origin of
Species, Joseph Reay Greene (1859) Professor of Natural
History in Queen’s College, Cork, Ireland provided a suc-
cinct introduction to the principles of systematic zoology,
including a clear account of what is meant by a natural
classification. This can be taken as a good example of the
post-Linnaean status quo in the mid-19th century, before
Darwin’s ideas began to firmly take hold. Greene thought
that the relations of living beings firstly concern “their
relations to one another” and secondly “their relations to
the conditions in which they are placed” (the “conditions”
being the external environment). For him, a natural classi-
fication results from an understanding of the former, while
the laws of geographical distribution may be deduced from
the latter. Specialised organs that correspond in structure
were classed as homologous (even if they differed in func-
tion). As such, they allowed for the determination of rela-
tions, whereas those that corresponded in function but not
in structure did not assist in this determination. Systems
which group animals simply by recognisable external
characters are for him “artificial” while he defines a true
“natural” classification as (with his own emphases):
… the right appreciation of the mutual relations of ani-
mals, as dependent on those characters and capacities that
they have received from their Creator. And as there is but
one Author of Nature, so also there can be but one true in-
terpretation of that Author’s plan … (Greene, 1859: xxiv)
Most importantly, Greene (1859: xxvii) finds little
positive evidence for the proposition:
28
TAXON 58 (1) • February 2009: 18–31
That one species may pass into another, or, in other words,
that species have no existence.… for it is more probable
that species should have been created with a certain degree
of variability, than that mutability should be a part of the
scheme of nature.
In short, despite the debate being fanned by Darwin
and his supporters, this mid-19th century view corresponds
closely with 18th century Wolffianism and creationism as
evidently accepted by Linnaeus via John Ray.
The prospect of Creative Evolution (organic evolution
as a product of God’s Creation) was indicated by Ray
(Pavord, 2005: 384) but was arguably first properly ar-
ticulated by Charles Darwin’s grandfather Dr Erasmus
Darwin (1731–1802). He did this through a debating group
of seminal thinkers and practical people (often dissident
protestants). From their sometimes wayward thinking and
because they met near each full moon, they were organ-
ised as the Lunar Society.
Erasmus Darwin (a friend of the mechanistic philoso-
pher Rosseau) was a physician, inventor and keen botanist.
He wrote Phytologia, or the Philosophy of Agriculture and
Gardening (1800) and philosophical poetry about plants,
e.g., The Botanic Garden (1791) with poems on the Economy
of Vegetation and The Loves of the Plants, an extravagant
versified tribute to the work of Linnaeus (Barber, 1980).
Erasmus Darwin and other members of the Lunar
group were certainly familiar with Linnaeus’ classification
and shared with him an enthusiasm for ordering ideas and
applying knowledge for practical and economic purposes.
However, the abstemious Erasmus Darwin evidently had a
negative, puritanical response to Linnaeus’ bawdy way of
expressing his sexual system for the classification of plants.
A disapproving attitude was maintained by many other
contemporary philosophers and scientists and persisted
into the 19th century (Barber, 1980: 54; Pavord, 2005: 398).
Indeed, this earlier public response may have, for a time,
inhibited Charles Darwin’s exposition of sexual selection
as a critical subset of natural selection in the origin of
species (Dixon & Brishammar, 2007). One Lunar Society
member James Watt (inventor of the steam engine) joked in
correspondence with Erasmus Darwin in 1781 that:
I do not know how steam engines come among the plants;
I cannot find them in the Systema Naturae, by which I
should conclude that they are neither plants, animals, nor
fossils, otherwise they could not have escaped the notice
of Linnaeus. However, if they belong to your system, no
matter about the Swede … (Pearson, 1943: 78)
Erasmus Darwin argued that:
Cause and effect may be considered as the progression,
or successive motions, of the parts of the great system of
TAXON 58 (1) • February 2009: 18–31
Nature.… This perpetual chain of causes and effects, whose
first link is riveted to the throne of God, divides itself into
innumerable diverging branches .… (Pearson, 1943: 127)
He published his ideas in Zoonomia, or the Laws of
Organic Life (1794) which is credited by the theologian
Samuel Butler and others as containing the first com-
plete, well-rounded view of evolution as a product of God’s
Creation. Pearson (1943: 136) states:
It was he and not his grandson Charles, who to quote Dr
Dowson was ‘the first to give a series of plausible reasons,
and the best that have yet been advanced, for believing the
Origin of Species by transmutation possible’.
On the contrary, Charles Darwin in his autobiography
of 1876 (published in 1929), expresses a disappointment
with the Zoonomia:
It has sometimes been said that the success of the Origin
proved that ‘the subject was in the air’ or ‘that men’s minds
were prepared for it’. I do not think that this is strictly true,
for I occasionally sounded not a few naturalists, and never
happened to come across a single one who seemed to doubt
about the permanence of species. (pp. 60, 61)
So it was left to Charles Darwin and Alfred Russell
Wallace to fully demonstrate that species had no such
God-given permanence or fixity and to propose natural
selection as an evolutionary mechanism. This admitted
the prospect of moving the theory of evolution beyond
theology and philosophy to scientific secularism.
Interestingly, Charles Darwin’s life and formative influ-
ences echo those of Linnaeus. Indeed, he declared himself to
be a Linnaean, was an active Fellow of the Linnean Society
of London and there delivered with Wallace the first (1858)
paper on the theory of natural selection. Both C. Darwin
and Linnaeus resisted family pressure to train for a life as a
minister in the protestant church. They engaged in medical
studies but eventually forsook this for wide-ranging research
in natural history, including expeditionary studies. Both rec-
ognised the importance of sex in biological explanation and,
in the face of public and theological criticisms, advocated the
recognition and acceptance of the close affinities between
humans and primates. They advanced their careers through
socially astute marriages and were deeply affected by the
early loss of their own offspring. To a great extent they were
sceptics or ‘free thinkers’, in Darwin’s case moving from
deism to agnosticism. Active and vigorous long-distance
travellers in their youth, Darwin and Linnaeus became dis-
tinctly preoccupied with personal health in their mature
years (Blunt, 1971: 228–229; Darwin, 1929). Surviving pe-
riods of public controversy, both were ultimately and rightly
applauded and honoured by civil and scientific society.
Reid • Linnaeus’ life and relationship to modern biology
In the great body of Linnaeus’ letters, manuscripts
and books are discernable foundations for Darwinian evo-
lution and many other later disciplines including: anthro-
pology, biogeography, bioinformatics, biomechanics, bio-
nomics, biotechnology, biological control, conservation,
ecology, epidemiology, ethnography, medical diagnostics,
microbiology, palaeontology, pharmacology, phylogeny
and the public understanding of science, education and
technology. Linnaeus’ contribution to modern science,
biology, medicine, systematic thinking and philosophy
has, therefore, been enormous, diverse and enduring.
ACKNOWLEDGEMENTS
In developing ideas for this paper I have had the great
advantage of discussions with distinguished Linnaean schol-
ars during the Tercentenary celebrations in Sweden, January
2007. I was invited to these as the Immediate Past President of
the Linnean Society of London. Lars-Ola Borglid was good
enough to provide a conducted tour of the key Linnaean sites
in Råshult, Växjö and Stenbrohult. Professor Carl-Olof Jacob-
sen, President of the Swedish Linnaean Society and former
Secretary General of The Royal Swedish Academy of Sciences
kindly introduced me to Linnaeus’ Uppsala and Hammarby. I
have also had the benefit of being generously hosted in Sweden
by Professor Mats Bergquist, Chancellor of Växjö University
and former Swedish Ambassador to London; by Professor An-
ders Hallberg, Vice Chancellor, University of Uppsala; by Eva
Bjorn, Curator Linnemuseet, Uppsala; and by Inger Lilequist,
Director-General, National Heritage Board of Sweden. I must
warmly thank Professor Tod Stuessy, University of Vienna, for
inviting me to give an address on Carolus Linnaeus at a Sym-
posium on Systematic Biology, at the 2007 Botanical Congress
in Chicago. This was organised with the unstinting support
of the American Society of Plant Taxonomists. That address
formed the basis of the present paper. During its preparation,
my good friend Gordon Howes (formerly of the Natural History
Museum, London) alerted me to an important but obscure work
of Professor Joseph Reay Greene (1859). Dr. James Dobreff,
Research Fellow at Lund University thoughtfully introduced me
to a project on Linnaeus’ lesser known apostle Daniel Rolander,
who wrote the under-appreciated Diarium Surinamicum. I am
most grateful to Gina Douglas acting Executive Secretary of the
Linnean Society of London and Lynda Brooks the Librarian for
patiently tracking down bibliographic references for me, includ-
ing in the Linnaean Correspondence archive. I must also thank
Dr. Polly Winsor for engaging in a stimulating correspondence
with me on typology and essentialism, points upon which we
may never agree! Professor Tod Stuessy, my wife Sally and
son Alex kindly criticised the manuscript for me and Claudine
Gibson helped with proof-checking. I am also greatly indebted
to two anonymous reviewers and the editors for their detailed
and helpful comments.
29
Reid • Linnaeus’ life and relationship to modern biology
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