494
Documenta Praehistorica XLVII (2020)
The biology of aesthetics
As a field of research, aesthetics generally does not
relate to biology or evolution, but primarily to cul-
ture and learning, although there is no doubt that
evolutionary theory can reveal a lot about aesthetic
preferences. Roughly speaking, aesthetics studies
what we like or dislike, and the important question
is whether standards of judgments can be learned or
are (to a greater or lesser extent) inherited. In line
with the tendency of social scientists throughout
most of the 20th century to ignore knowledge that
came from biology (Degler 1991), theorists of aesthe-
tics and the arts mostly did not advocate ideas about
the existence of human nature, but emphasized his-
toricist explanations based on historical contexts
and cultural factors. Thus, aesthetic values are usu-
ally seen as referring to the prevalent social values
in a particular culture, and that kind of social or cul-
tural constructivism also implies relativism regard-
ing aesthetic values.
The situation changed with the rise of evolutionary
psychology, which began to advocate aesthetic uni-
versalism (e.g., Dutton 2003; Thornhill 1998; 2003;
Dissanayake 1992), and the answer to the question
of why the position of universalism is compatible
with evolutionary psychology is simple. All organi-
Habitat selection and the evolutionary aesthetics
of landscape preference
Marko {korić, Aleksej Ki[juhas
Faculty of Philosophy, University of Novi Sad, Novi Sad, RS
mskoric@ff.uns.ac.rs
aleksej.kisjuhas@ff.uns.ac.rs
ABSTRACT – This paper analyses the processes of habitat selection and human landscape preferences
from an evolutionary perspective, with the aim of demonstrating how humans aesthetically choose,
assess and aspire to live in an environment in which our species and our ancestors evolved in dur-
ing the pre-Neolithic period. We present the basics of evolutionary aesthetics, then analyse the pro-
cess of habitat selection and the most influential evolutionary theories of landscape preference.
Finally, we refer to applied empirical research and point out that a comprehensive evolutionary the-
ory must also take into account the psychological and cultural elements that affect human well-being.
IZVLE∞EK – V ≠lanku analiziramo proces izbire ∫ivljenjskega prostora in preference ljudi pri izbiri
krajine iz evolucijske perspektive z namenom, da poka∫emo, kako ljudje na podlagi estetike izbira-
jo, ocenjujejo in stremijo k ∫ivljenju v okolju, v katerem so se v obdobju pred neolitikom razvijali
na∏a vrsta in na∏i predniki. Predstavljamo osnove evolucijske estetike, nato analiziramo proces iz-
bire ∫ivljenjskega prostora in najbolj vplivne evolucijske teorije o krajinskih preferencah. Na koncu
se sklicujemo na uporabne empiri≠ne raziskave in poka∫emo, da mora celovita evolucijska teorija
upo∏tevati tudi psiholo∏ke in kulturne elemente, ki vplivajo na dobro po≠utje ljudi.
KEY WORDS – evolutionary aesthetics; habitat selection; biophilia; landscape preference
KLJU∞NE BESEDE – evolucijska estetika; izbira ∫ivljenjskega prostora; krajinske preference
Izbira /ivljenjskega prostora
in evolucijska estetika krajinskih preferenc
DOI> 10.4312\dp.47.28
Habitat selection and the evolutionary aesthetics of landscape preference
495
across cultures, but research shows that some basic
algorithms are universal and have evolutionary ori-
gins (e.g., Brown 1991). Like physical characteristics,
our cognitive structures are adaptations to solving
the reproductive problems of the human phyloge-
netic past. This implies that evolutionary aesthetics
is part of evolutionary psychology and rests on the
premise that human basic aesthetic preferences
have evolved in order to improve survival and re-
productive success. In other words, evolutionary
aesthetics is an attempt to understand the aesthetic
reasoning of humans and their spontaneous differ-
entiation between ‘beautiful’ and ‘ugly’ as a biologi-
cally adaptive capacity for making important deci-
sions in life (see also Averill et al. 1998; Dutton
2003; Hartmann, Apaolaza-Ibanez 2010; Paden
et al. 2012; Seghers 2015).
Probably the most famous example of evolutionary
aesthetics is physical attraction in the context of
mate selection, of which much has been discussed
(e.g., Symons 1979), and the same logic can be ap-
plied to other visual preferences. In this paper, the
emphasis is on landscapes and habitat selection pre-
ferences. The basic premise of evolutionary aesthe-
tics in this domain is to show how people choose and
want to reside in an environment where our species
and ancestors evolved for millions of years.
We can differentiate among direct, indirect and vi-
carious (or symbolic) experiences of nature. It is
obvious that in the modern world there has been
a decline in direct experience of the natural environ-
ment, as it involves spontaneous contact with na-
ture. Indirect experience represents organized con-
tact with nature, where human intervention is seen
(e.g., zoos, parks), and symbolic does not mean
actual but mediated contact (books, movies, etc.).
Thus, the growth in indirect and symbolic experience
is noticeable and direct contact with nature has
fallen dramatically. So the important question is
whether and to what extent contact with natural
systems and processes is important and useful for
the well-being of today’s humans, living in an envi-
ronment that does not resemble the one in which
our ancestors evolved.
Habitat selection
Humans receive a wealth of information from their
surroundings on a daily basis, and it is impossible to
assimilate and use all of it, while it is clear that most
of this information has very little or no value to our
survival. That is why natural selection has made us
sms are the products of evolution, which is not
brought into question by scientists, while there is
some controversy concerning the impact of evolu-
tion on behaviour (e.g., Segerstråle 2000). Evolutio-
nary psychologists acknowledge the influence of
evolution through natural selection both on the be-
haviour and mind of humans, and according to
these ideas human nature was formed long before
the Holocene, and endows us with certain capacities
which are not consequences of socialization, learn-
ing, traditions and the like (e.g., Tooby, Cosmides
1992).
We argue that there is sufficient convincing evidence
to conclude that human nature exists in the form of
genetic biases that affect our perception, decision-
making and behaviour in general (e.g., Wilson
1978). Cultural evolution is influenced by biology,
although the biological evolution of the brain has
taken place in a social context, so it is not disputed
that culture plays a large role in human behaviour.
Human culture is part of human biology, thus any at-
tempt to separate them is artificial and misguided
(e.g., Boyd, Richerson 2005). Only from an evolu-
tionary perspective can we understand why our
knowledge fits our environment; that is, how we
manage to understand the world around us. Like-
wise, an evolutionary perspective offers an explana-
tion as to why we know so much about the world,
although we have very limited personal experience.
And finally, only through evolution can we explain
why our knowledge reflects the environment in
which our ancestors evolved (e.g., Campbell 1974).
Due to evolution, our perception and cognition are
selective, and we can speak of perceptual biases that
occur at three different levels: basic biases, which
we share with higher vertebrates, biases that are
characteristic of our species, and specific cultural
perceptual biases. Similarly, our emotions have
evolved since they have positively affected the sur-
vival and reproductive success of our ancestors (To-
oby, Cosmides 1990), indicating that humans res-
pond emotionally to the world around them because
they possess an aesthetic sense which is the product
of evolution through natural selection. Studying both
cognitive and emotional preferences can thus serve
as a bridge between psychology and evolutionary
theory (Kaplan 1987).
The universality of art in all known cultures indicates
that aesthetics is also closely related to psychologi-
cal adaptations (Tooby, Cosmides 2001). Aesthetic
reasoning undoubtedly varies greatly over time and
Marko {korić, Aleksej Ki[juhas
496
develop neural programs which assess information
and act as filters that allow important data in and
prevent irrelevant data from entering. These filters
constitute biologically prepared learning that pays
particular attention to decisions that affect survival
and reproduction, which is why being attracted to
certain environmental information is adaptive be-
haviour (Kaplan 1987). However, these decisions
do not need to be adaptive today, since natural se-
lection cannot affect our fit with future environ-
ments. For example, phobias that are almost univer-
sal in humans relate to snakes, spiders, heights, dark-
ness and the like, all of which have had a major im-
pact on survival and reproduction during most of
human evolution, and not with the knives, automo-
biles, guns, and so on which are really more dange-
rous to human life in modern environments. As such,
many phobias have to do with the environment in
which human nature was formed, and they have ma-
naged to survive since then.
It is reasonable to assume that habitat selection is a
process associated with human survival; that is, which
has left consequences for the survival and reproduc-
tion of humans and their ancestors. Until the Neoli-
thic and urban development, people mostly lived a
nomadic and hunter-gatherer lifestyle, and the search
and selection of habitats were of utmost importance
not only for providing food and shelter, but also be-
cause habitats evoke certain emotions, stimulate en-
vironmental exploration, give sense of well-being
and identity, etc. (Hunziker et al. 2007). Simply put,
individuals who were able to find and identify a ha-
bitat that offered protection from predators and bad
weather, while also enabling access to food, water,
and other resources, were more successful than in-
dividuals who could not recognize such qualities in
a given environment (Ruso et al. 2003). Of course,
the preference for particular habitat types that fa-
cilitate adaptation is not unique to the human spe-
cies alone, so habitat selection is widespread among
vertebrates, and has been reported even in animals
raised in the laboratory and without any prior expe-
rience with the natural environment (Kaplan 1987;
see also Wecker 1964). Therefore, it can be said that
habitat selection is an almost universal activity among
animals that affects many choices of individual orga-
nisms.
Environmental preferences co-evolve with its quali-
ties, meaning that organisms respond positively to
environments where the chances of their survival
and reproductive success are good (Orians, Witten-
berger 1991). Likewise, organisms that choose to
settle in less conducive environments leave fewer
offspring. However, this correlation is not so simple,
because a habitat which seems good at first glance
may be inadequate due to other factors (such as in-
fectious diseases, hidden predators, etc.), and orga-
nisms may have difficulty in truly, accurately or pre-
cisely assessing the quality of an environment (Ori-
ans, Wittenberger 1991). The habitat selection pro-
cess itself takes place in stages – encounter, explore
and exit or establish – and thus is a kind of hierar-
chical process that involves making several strategi-
cally important decisions.
The first stage begins with the arrival of the organ-
ism in an unknown general area, where there are se-
veral habitats that are suitable for various activities
– some are for courtship, some for hunting, settling,
etc. The organism must first decide whether or not
a certain habitat should be explored or whether it
should move on. This decision is largely based on
the judgment of the distribution of objects in space,
water, trees, or vegetation, shelter, potential distance
from prey or a safe place, and the like. If an organ-
ism considers staying, then it goes into a more de-
tailed exploration of the environment and then de-
cides whether to stay or not, as well as whether that
possible stay will be shorter or longer (Orians, Wit-
tenberger 1991).
It is also clear that the key factor in this process is
time, since better habitats are usually already occu-
pied, which is why organisms have to make deci-
sions very quickly – almost instinctively and often
based on incomplete information. Contemporary ha-
bitat selection research shows that subjects do ex-
press their preferences relatively quickly and easily,
but most often cannot explain their choices and are
unaware of the predictive variables that make them
prefer one environment rather than another, imply-
ing some automaticity in these choices (Kaplan
1987).
From an evolutionary perspective, there are many
adaptive benefits to quick, automatic judgment of
the information traits concerning a place or space
we are approaching. Walking on varied terrain, cho-
osing the right path, etc., require constant re-evalu-
ation as the landscape opens up new vistas and op-
portunities. It is the information processing speed
that is crucial for rapid response, which makes it ap-
propriate to be automatic and unconscious, and to
result in an affective (rather than cognitive) reaction
(Ulrich 1983). In other words, it is not only impor-
tant to rationally identify habitats that are useful,
Habitat selection and the evolutionary aesthetics of landscape preference
497
but also environments that we like emotionally. It
is an effective guide to ongoing behaviour when an
individual is intuitively (and aesthetically) attracted
to promising places, or rejects unpromising ones.
Although today habitat selection is not a particular-
ly important factor in human survival, this process
was extremely significant for the everyday survival
of our ancestors (Ruso et al. 2003). Therefore, the
psychological mechanisms underlying habitat selec-
tion have been the subject of intense selection pre-
ssures, including the emergence of emotional reac-
tions to certain environmental characteristics (Kap-
lan 1987; Seghers 2015). Because habitat selection
has been crucial throughout human evolutionary hi-
story, research into this phenomenon, especially in
the context of landscape preference, is of great im-
portance for the general understanding of our evolv-
ed aesthetic tastes (Ruso et al. 2003).
Biophilia and the evolutionary theories of land-
scape preference
Roughly speaking, two basic types of evolutionary
aesthetics can be distinguished – evolutionary intu-
itionism and evolutionary cognitivism (Paden et al.
2012). Intuitionism speaks of fundamental aesthetic
intuitions as adaptive preferences, meaning that the
pleasures underlying aesthetic reasoning evolved
because they played a role in adapting human beings
to their environment. On the other hand, evolutio-
nary cognitivism is based on remarks about the in-
trinsic beauty of (certain) living beings or landscapes;
that is, the view that evolution brings about beau-
tiful living forms, remarkable natural phenomena,
and so on. Likewise, there are two paradigms when
it comes to landscape preferences – objectivist and
subjectivist (Lothian 1999; Maulan et al. 2006).
Objectivist approaches perceive visual quality as in-
herent in the landscape, while subjectivists claim
that visual quality is a construct of the beholder.
Within subjectivist approaches, positivist and phe-
nomenological models exist – positivists analyse the
physical and measurable features of landscapes,
while phenomenological models focus on personal
experience and meaning (Ohta 2001; Thwaites,
Simkins 2007). It is just about what we attach the
greater importance to, because it is indisputable that
the landscape quality depends on both objective and
subjective factors (Daniel, Vining 1983).
Thus, objectivists usually assume that there is a cer-
tain unchanging standard for aesthetic appreciation
in the characteristics of the object itself (line, colour,
form, etc.), and its proponents are most often artists,
(landscape) architects, some physical geographers,
ecologists and environmentalists, and others who
claim aesthetic qualities of the object rest in its for-
mal qualities independent of human perception and
interpretation. Many of them also insist that aesthe-
tic quality stems from ‘naturalness’, i.e. ecological di-
versity, and that unmodified or ‘natural’ landscapes
are therefore of greater aesthetic value to humans.
On these theoretical foundations, and the concept of
‘natural’ and ‘picturesque’, for example, Hyde Park
in London was designed, with its curvilinear lines,
vegetation masses and open vistas (Maulan et al.
2006).
On the other hand, subjectivist approaches empha-
size that landscape aesthetics are the result of the in-
teraction between the observer and object, which is
why they focus on the cognitive and affective reac-
tions of individuals. In this sense, the aesthetics of
the landscape is a human construct on the socio-cul-
tural or individual level, since it rests on the percep-
tion and interpretation by the human mind. In plain
language, ‘beauty is in the eye of the beholder’. It
can be said that the objectivist paradigm often pla-
ces people in a relatively peripheral position in
which they, similar to a camera, passively ‘capture’
landscapes (like a work of art), while the subjecti-
vist paradigm also takes into account the variables
of human knowledge, experience, emotions, needs,
etc. (Maulan et al. 2006.29–30).
The evolutionary theories that interest us in this pa-
per imply that beauty or ugliness are not intrinsic
features of objects, but that they arise from the in-
teraction between features of objects and the hu-
man nervous system. This indicates that for humans,
beautiful objects are those that (accompanied by a
positive reaction) improve our lives, in terms of in-
creasing the chances of survival and reproduction.
In contrast, ugly ones are those that impede certain
aspect of our lives (Tooby, Cosmides 2001). If aes-
thetic preferences are the product of evolution, it
follows that they are adaptations to the Pleistocene
environment and not necessarily to the contempo-
rary one, since the social environment in which we
live today has existed for an extremely short time
when viewed in the context of the evolution of the
human species.
Therefore, there are Darwinist-oriented theories
which explain landscape preferences as being the re-
sult of human evolution, implying that what we con-
Marko {korić, Aleksej Ki[juhas
498
sider beautiful today is something that increased our
ancestors’ chances of survival. For this reason, even
today, a great deal of the everyday aesthetic experi-
ence of humans involves a cognitive, emotional and
behavioural response to landscapes in the immediate
environment (Russo et al. 2003). In the following
passages we will present the most significant evolu-
tionary theories of landscape preferences and aesthe-
tics, such as the prospect-refuge theory, savanna or
habitat theory, information processing theory (or the
mystery and complexity theory) and affective theo-
ry. They can all be linked to a more general frame-
work called the biophilia hypothesis.
This hypothesis mainly relates to the claim that hu-
mans have a basic need to interact with ‘nature’
(Averill et al. 1998). As early as in his 1963 essay,
Erich Fromm (Fromm 1963; Eckardt 1992) made a
distinction between necrophilia (attraction to death
or love of death) and biophilia (attraction to life or
love of life). Fromm also mentions the biophilic per-
sonality type, which is typical of individuals who are
surrounded by people who love life, security, justice
and freedom. In a different sense, the term was re-
vived and popularized by Edward O. Wilson (1984;
Kellert, Wilson 1993) who defines it as: “an innate
tendency to focus on life and lifelike processes”
(Wilson 1984.1), or an “innately emotional affilia-
tion of human beings to other living organisms”
(Wilson 1993.31). This would suggest that humans
depend on nature, but not just for mere survival –
there are also aesthetic, intellectual and cognitive
reasons why this relationship is significant. The es-
sence of this idea is to explain the human biological
need to maintain a relationship with life and lifelike
processes, which is biologically based, so it repre-
sents the evolutionary heritage of our species.
Similar ideas can be found in the field of ecopsycho-
logy, which discusses ecological unconscious (Roszak
1992), and whose main purpose is to restore the con-
nection between humans and nonhuman nature. It
refers to the sense of interconnectedness between
humans and other living beings, which stems from
our ancient ancestors. Modern life usually prevents
most people from understanding this connection,
but Roszak points out that there is environmental
reciprocity and that in some way we sympathize
with planet Earth.
In the context of this paper, the most important
question is whether human physical and mental
well-being depends on contact with natural systems
and processes (or not), primarily because humans
throughout their history lived in environments that
were not altered to the extent those they are living
in today are altered. Usually, the social dimension of
the world is cited as the key environment in which
humans evolved, and while no one denies the im-
portance of social interactions and sociality (πkori≤,
Ki∏juhas 2015), the importance of vegetation, land-
scape, the living world (plants and animals), wind,
rain, smells, sounds, etc., must not be omitted. This
is not to say that biophilia, like an instinct, is rigidly
encoded in our genome, but a kind of weak biologi-
cal tendency, a set of predispositions for gaining cer-
tain preferences. It is simply a part of human nature
that was formed in an evolutionary environment
(including the physical environment) through biocul-
tural evolution, and therefore depends on learning,
experience and culture (Boyd, Richerson 2005).
When it comes to landscape preferences, probably
the most important question is the one concerning
the cognitive mechanisms involved in the process.
The most commonly mentioned mechanism is sim-
ilar to the ethological construct of the innate sche-
mata, which is a mental image of the ideal landscape
that represents the standard of judgment. Theoreti-
cal differences generally exist when it comes to the
essence of that image – be it a landscape in which
humans evolved (such as savanna), or perhaps any
type of landscape containing those features that pro-
mote fitness. In this sense, among the first evolutio-
nary theories of landscape preference that stands
out is what was then called habitat theory by Jay Ap-
pleton (1975), now known as prospect-refuge theo-
ry. He was one of the earliest to suggest the provo-
cative idea that a preference for a particular type of
landscape is part of our evolved heritage, and that
the environment should be viewed in functional ra-
ther than morphological frames. Beauty (landscape)
is not found neither in beautiful objects, nor in the
eyes of the beholder, but in the (functional) relation-
ship between the individual and environment.
In this regard, aesthetic satisfaction comes from the
fact that the observer sees an environment as beau-
tiful or useful for fulfilling his or hers biological ne-
eds. The possibility of “seeing and not being seen”
satisfies many such needs, as previously written by
Konrad Lorenz (1949/1952). Therefore, Appleton
claims that the key components of landscape prefe-
rences are ‘prospect’ (having a grand view, over-
view or opportunity) and ‘refuge’ (having a safe
place to hide or safety). For example, closed forests
are not good for prospect and deserts are not good
for refuge, while savannas offer a good combination
Habitat selection and the evolutionary aesthetics of landscape preference
499
of these elements. Aesthetically most attractive and
pleasing are landscapes that have a balance between
prospect and refuge components – elevated land-
forms (due to the overall view of the landscape in
search of food, water and prey) and attributes such
as groupings of trees (due to safety factors), with
open spaces and fresh water (see also Maulan et al.
2006). Appleton argues that such evolved preferen-
ces apply today, regardless of the fact that most peo-
ple no longer have to seek shelter from predators.
However, it can be said that his theory deals only
with a limited part of the otherwise very complex
judgments that people make about their environ-
ment (Heerwagen, Orians 1993). Appleton looks
only at the initial evaluation and exploration of un-
familiar environments and focuses on the opportuni-
ties to acquire information, as well as the security
that can(not) be achieved, while the habitat selec-
tion process is usually much more complex. Some
empirical tests confirm his theory (e.g., Clamp, Po-
well 1982; Mealey, Theis 1995) and other do not
(e.g., Klopp, Mealey 1998). There are not many stu-
dies examining the balance between prospect and
refuge, since it is very difficult to analyse – for exam-
ple, it is not clear whether darkness counts as pros-
pect or refuge (Bunkse 1977), and this theory is in-
deed quite reductionist and extreme (Ulrich 1983).
That is why Appleton later (Appleton 1990) tried to
modify the theory, although to date there is still not
much convincing evidence to support it.
The next influential evolutionary theory is common-
ly called the savanna hypothesis or savanna theory,
and it also rests on assumptions concerning the
search for a suitable habitat (Orians 1980; 1986;
Orians, Heerwagen 1992). Gordon Orians argues
that humans have an innate preference for environ-
ments that are similar to the African savannas, be-
cause it was not only the habitat of our distant an-
cestors, but also the spatial context of hominid evo-
lution, which is why they evolved a preference for
environments of similar appearance. At the same
time, it is argued that the (evolved) preference for
savannas (i.e. environments containing grasslands
and scattered trees with water nearby) offered an
evolutionary advantage to hunter-gatherers, which
is complementary to the prospect-refuge theory (Ap-
pleton 1975).
These open landscapes also offered benefits for ex-
ploring the environment, as well as the highest con-
centration of resources for early humans and species
before them. This microenvironment allowed hu-
mans to survive because they had food, a resting
place, shelter from predators, sun shade and nearby
animals that could be hunted (see also Seghers
2015). Some studies show that in many countries
around the world people arrange their landscapes to
resemble natural savanna landscapes, i.e. as a mix
of open grassland and groups of trees, just as sa-
vanna-like landscapes appear on many paintings, ca-
lendars and (desktop) wallpapers (Orians 1980; see
also Heerwagen, Orians 1993; Hunziker et al.
2007).
However, this theory also has its critics, who claim
that human ancestors began to settle in non-savanna
environments millions of years ago, which actually
left them plenty of time to evolve aesthetic preferen-
ces toward different environments too (Diamond
1993). When it comes to more empirical research,
some authors offer limited support for the human
savanna tendency, which has been confirmed in
children between the ages of eight and eleven (Bal-
ling, Falk 1982), while other research has not con-
firmed this hypothesis (e.g., Lyons 1983).
The third relevant evolutionary theory of landscape
preference is the one that asserts factors such as
mystery and complexity, or the so-called informa-
tion processing theory (Kaplan 1987; Kaplan, Kap-
lan 1989). Like Appletone, Stephen and Rachel Kap-
lan point to two basic human needs that influence
aesthetic appreciation of landscape – the need for
exploration and for understanding. They claim that
human ancestors who depended on hunting and ga-
thering had to explore new areas, but also to under-
stand them. This is why these authors apply the in-
formation processing approach to landscape aesthe-
tics in order to explain human-landscape interac-
tions, stating that people are trying (and striving) to
make sense of the environment through four pre-
dictor variables: (1) coherence (immediate under-
standing of the fit and harmony of elements in the
environment), (2) complexity (visual wealth that can
be directly explored), (3) legibility (understanding
of what we see and the ability to find a way and not
to get lost), and (4) mystery (the existence of new
things to explore if we go deeper and further into
the landscape). Two variables help us understand
the environment (coherence and legibility), while
the other two (complexity and mystery) inspire us
to explore it and influence our (landscape) aesthetic
preferences.
For example, there is some regularity in preferences,
even when it comes to nature scenes in photographs
Marko {korić, Aleksej Ki[juhas
500
(Kaplan et al. 1972). The most positively evaluated
scenes were those that “(1) contained a trail that
disappeared around the bend, (2) depicted a
brightly lit clearing, partially obscured from view
by intervening foliage” (Kaplan 1987.8), suggesting
that the most popular images ‘promise’ more infor-
mation. For these reasons, this predictor is also called
‘mystery’, while some characteristics such as coher-
ence or complexity are much less significant. The ex-
planation for this phenomenon is based on a claim
concerning the adaptive function of human curios-
ity – humans prefer those environments that facili-
tate the acquisition of new information, i.e. those
where new information can be obtained if the envi-
ronment is explored.
Mystery, then, is primarily about the expectation that
a deeper encroachment into the landscape enables
access to additional and new information, as a bal-
ance between what is seen and what is anticipated.
It is assumed that the survival of the human species
depended on the development of cognitive informa-
tion processing skills that have further developed in-
to a preference for the landscapes which made sense
for humans. Joachim Wohlwill also explored the as-
sessment of non-representational art, urban and ru-
ral scenes, and natural landscapes with an emphasis
on elements of complexity (colours, shapes, textu-
res, etc.) (Wohlwill 1968), and the data indicated
that individuals mostly preferred images of mode-
rate complexity.
Finally, the affective theory by Roger Ulrich (1977;
1983; 1986) claims that natural environments and
landscapes give rise to emotional states of well-being
in humans that can be measured. He also analysed
the information features of different landscapes and
identified the following five significant variables: fo-
cality (coherence, unity), ground surface texture,
depth, mystery and complexity. However, Ulrich no-
tes that affective reactions to certain visual configu-
rations of these variables have had adaptive value
during human evolution, which is why people still
prefer them today, even without cultural learning or
socialization. The important emotions in this regard
are pleasantness, calm, exhilaration, caution, fear
and anxiety. It is a model that contrasts with Kap-
lans’ cognitive theory, because it is based on the idea
that emotional reactions to landscapes occur before
cognitive information processing (Zajonc 1980), as
well as the thesis that positive emotions have a be-
neficial effect on survival. In addition, Ulrich further
(Ulrich 1979; 1984; 1993) investigated emotional
responses to urban and natural landscapes, where
the results suggested that urban scenes evoke more
negative emotions.
Of course, in addition to evolutionary theories, there
are numerous cultural theories that explain land-
scape preferences through the process of learning
and influence of social, cultural and personal fac-
tors. In other words, aesthetic preferences are un-
derstood as social constructs, as it is thought that hu-
mans have repeatedly adapted to different habitats
and are very flexible when it comes to responding
to them. Here, the cognitive assessment of the func-
tions offered to individuals by a landscape is empha-
sized, rather than the immediate affective response
(Bell 1999).
The best known are the cultural theories highlight-
ing topophilia and ecological aesthetics. Topophilia
refers to the fact that people tend to bond with what
they know well, which means landscape preference
is influenced primarily by familiarity and experience
(Tuan 1974). The second group of theories emphasi-
zes the importance of knowledge about ecological
functions, and that knowledge leads to the preferen-
ces for a landscape (Carlson 2009; Gobster 1999).
Another group of theories speaks of genius loci
(meaning a specific atmosphere of a place), and thus
the unique and visually striking features and charac-
teristics of landscapes (Bell 1999). Still another
group of cultural theories notes the importance of
landscape heritage, where there are visual cues of
cultural heritage (Fairclough et al. 1999), while the
aesthetics of care foregrounds the importance of
signs indicating some concern about the landscape
(Nassauer 1992). There is not necessarily a contra-
diction or a discrepancy between evolutionary and
cultural theories, as evidenced by the conclusions of
some empirical research.
Applied empirical research and biophilic de-
sign
Numerous empirical studies show that aesthetically
beautiful and attractive habitats for humans are the
ones in which the evolution of our ancestors took
place, and children also mainly favour savannas,
landscapes with clouds and water, open spaces with
trees that fork near the ground, which offer low-
hanging fruits and the like. For most people, run-
ning water is more attractive than stagnant water,
remote mountains are more attractive than flat ter-
rain, and the most preferred are landscapes that are
relatively open and smooth with wide horizons, low
and homogeneous vegetation and scattered trees
Habitat selection and the evolutionary aesthetics of landscape preference
501
(with a round shape), clouds, game animals and the
like (Calvin et al. 1972; Kaplan et al. 1972; Balling,
Falk 1982; Kaplan, Kaplan 1989; Orians, Heerwa-
gen 1992; Purcell et al. 1994; Summit, Sommer
1999; Hartmann, Apaolaza-Ibanez 2010; Dutton
2003; Thornhilll 1998; 2003).
For example, in perhaps the most famous empirical
study of this type (Balling, Falk 1982), a sample
consisting of 548 very different respondents (ele-
mentary school students, college students, the elder-
ly, professional foresters, biology teachers, etc.) were
shown five different biomes on slides (tropical for-
est, desert, savanna, deciduous and coniferous forest,
and without images of water, animals, humans or
human intervention), and subjects under the age of
15 clearly displayed a preference for savanna-like
environments. For these reasons, it can be said that
there is limited support for the savanna hypothesis
(Orians 1980, 1986), due to the long evolutionary
history of human life in the savanna, which is best
expressed in childhood. The preference for savan-
nas decreases with age and experience, and a pref-
erence for the most familiar environments increas-
es (Balling, Falk 1982.25). This suggests that famili-
arity in terms of growing up and living close to a
(different) natural environment probably modifies
the initial preferences for savannas (Lyons 1983).
On the other hand, in a recent study of 750 subjects
and with images of 13 different biomes (savanna
trees, Canadian trees and lakes, birch trees in Eu-
rope, eucalyptus trees in Australia, an urban envi-
ronment, desert, etc.), the hypothesis of an innate
human propensity for savannas was not corrobo-
rated, but it was confirmed for lush green land-
scapes with a source of water and generally familiar
biomes (Hartmann, Apaolaza-Ibanez 2010). These
researchers therefore conclude that while an evolu-
tionary approach to landscape preferences is impor-
tant, limiting it to hunter-gatherer periods in the Pa-
laeolithic or the African savanna probably does not
capture all the dynamics of human behavioural evo-
lution. However, the mentioned familiarity effect was
confirmed (see also Herzog et al. 2000), where cen-
tral European mountains or the Mediterranean coast
are preferred over, for example, Australian eucalyp-
tus shrubs. The concrete application of the results
of this research concerns the domain of advertising,
that is, the claim of a more positive influence of
those commercials representing ‘natural’ scenes
with the favoured biosphere, than those with, for
example, urban environments or deserts without ve-
getation.
In an effort to demonstrate the savanna hypothesis,
however, additional research has been conducted
among individuals living in the rainforest belt of Ni-
geria (Falk, Balling 2010). After presenting five bio-
mes (rainforest, deciduous forest, evergreen forest,
desert and savanna), these subjects selected the sa-
vanna as the most suitable place to live. In addition,
the analysis of particular historical patterns of land-
scape design (private gardens, urban parks, etc.) in-
dicates certain universal characteristics or parallels,
such as savanna-like combinations of short grass and
scattered trees (Falk, Balling 2010.479). The results
of this study suggest that humans, regardless of their
background and experience, are likely to start life
with a preference for savanna-like environments.
Similarly, in a cross-cultural study of tree preferen-
ces (Orians, Heerwagen 1992), the respondents se-
lected as the most attractive trees the ones most re-
sembling trees from the East African savanna. Speci-
fically, people prefer trees with lower trunks (easier
to climb), with moderate canopy density (allowing
a balance between hiding and not being able to see
out), with high degrees of canopy layering (greater
opportunities for viewing out) and a broader tree
canopy relative to its height (because it can accom-
modate more people on branches). Large canopies,
short trunks or acacia trees, typical of the African sa-
vanna, are also preferred over oak, conifers, palm
trees or eucalyptus (Summit, Sommer 1999). A de-
tailed analysis of landscape features in Western
painting (Heerwagen, Orians 1993) also demon-
strated a human propensity for savanna-like envi-
ronments.
Theoretical and empirical knowledge of landscape
preferences can be applied in the context of envi-
ronmental education (Swonke 2000). Specifically,
biophilic feelings, as well as aesthetic preferences
for certain elements of the natural environment, can
serve to nourish the desire for environmental pro-
tection, that is, the development of the conservation
ethic (Wilson 1993). However, it is important to un-
derstand that mere contact or direct experience with
nature does not automatically lead to biophilia or
‘love’ for nature, and especially not to the preserva-
tion of the natural environment (Swonke 2000).
Some authors have empirically investigated the hu-
man tendency for fractals, i.e. those geometric sha-
pes that can be divided into parts that simultaneous-
ly resemble the original shape (Hagerhall et al.
2004). Fractals are relatively common in nature and
these authors claim that the most preferred are land-
scapes in the mid-range (of complexity) related to
Marko {korić, Aleksej Ki[juhas
502
fractal dimension, which are landscapes that also
correspond to savannas.
However, perhaps the most important application of
this knowledge concerns human health. Research
shows that preferred landscapes arouse positive
emotions and that even the pulse rate decreases
when observing natural (but not urban) landscapes
(Laumann et al. 2003). Furthermore, after surgery
patients who have a view of a natural landscape re-
cover better and faster (Ulrich 1983; 1984; Fuller et
al. 2007; Grahn, Stigsdotter 2003; Kaplan 1995;
Parsons et al. 1998; Rappe, Kivelä 2005).
Considering the evolutionary aesthetics and theories
of landscape preferences, highly illustrative is the
project The People’s Choice, realized by the artists
Vitaly Komar and Alexander Melamid in 1993 (Wypi-
jewski 1997). In collaboration with opinion polling
agencies, and while deliberately avoiding the word
‘art’, they explored the aesthetic preferences of peo-
ple from eleven countries from different continents.
It was meant in part as an artistically ironic commen-
tary on popular tastes, but also on political systems
that rely too heavily on polls and popular opinion
surveys when making decisions (Hillings 1999).
It turned out that the respondents’ favourite colour
was blue and their second favourite was green, that
people preferred realistic images depicting water,
trees and other plants, clouds, people (primarily wo-
men, children and historical figures) and animals (es-
pecially large mammals, domestic and wild), which
is highly consistent with the evolutionary theories of
(landscape) aesthetic preferences (Dutton 2003). Fi-
nally, it is particularly interesting that, based on
these popular responses, the artists
Komar and Melamid actually painted
the image that incorporated all the
highest-rated elements from their re-
search for each country.
In the case of the United States, it
was (a rather ‘kitschy’) blue-green
19th century realist landscape, featur-
ing children, deer, and George Wa-
shington (Fig. 1). With certain cultu-
ral-specific exceptions (e.g., Russians
chose Jesus Christ instead of George
Washington as their favoured histori-
cal figure), a very similar pattern was
observed in other societies, which is
why the authors conclude that it is a
“universal model for the paintings
that people want” (Hillings 1999.60). In a more con-
temporary context, it is also interesting to refer to
the famous default computer desktop wallpaper for
the Windows XP operating system, a photograph of
a green hill and blue sky with clouds (from Sonoma
County, California), conspicuously named ‘Bliss’, and
which can be considered as one of the most viewed
photographs in the 2000s (Fig. 2). On the other
hand, this extremely popular image still lacks people,
animals, and especially trees (in the sense of refuge).
Some research (e.g., Kaplan et al. 1972) clearly
shows that people often prefer images of the natu-
ral over the built environment, and that out of the
latter the most attractive image is that of an urban
park, the lowest ranked of the ‘natural’ images. Bio-
philic design or architecture is built on these bases,
and it emphasizes the importance of one’s experi-
ence of nature in the built environment which takes
into account the ethics of environmental sustainabil-
ity (Söderlund 2019). In other words, it is an at-
tempt to reintegrate people with the natural environ-
ment after a long time: “the expression of the in-
herent human need to affiliate with nature in the
design of the built environment” (Kellert, Heerwa-
gen 2008.viii). There is also the idea of biophilic ci-
ties, which are not the same as green cities, since the
mere presence of nature is not enough. In biophilic
cities, people are directly and actively involved in
various processes, such as learning about nature, en-
joying it or taking care of it, and developing emo-
tional bonds towards it (Beatley 2011; Kellert et al.
2008).
Biophilic design can thus be organic and vernacular.
Organic design is based on theory and research on
Fig. 1. Komar and Melamid, “America’s Most Wanted”, 1994. Oil
and acrylic on canvas.
Habitat selection and the evolutionary aesthetics of landscape preference
503
how humans respond positively to natural materi-
als, lighting and shapes, and the main goal of bio-
philic architecture is to maximize well-being while
minimizing ecological impact. Vernacular design, on
the other hand, aims to awaken a sense of connec-
tion to place (see also Hunziker et al. 2007), which
at the same time might create a desire within a cer-
tain community to care for a particular bioregion.
Environmental psychology also speaks of the ten-
dency of people to create and develop a kind of at-
tachment with specific locations. This attachment
can then relate to our immediate environment, such
as our home, but also to the neighbourhood, city or
even the region. However, place attachment does not
necessarily have to do with behaviour that takes care
of nature in general, so in addition to the undoubted
importance of the environment for survival, evolu-
tionary theories must also take into account the emo-
tional bonds between people and place. There is no
doubt that place can be meaningful for humans, and
that the aesthetics of survival is not enough to fully
explain one’s environmental preferences.
Why the aesthetics of survival is not enough
Philosophical aesthetics is not threatened by an un-
justified reduction to evolutionary biology, psycho-
logy or neurobiological explanations, while evolu-
tionary disciplines and humanities may be comple-
mentary in understanding the human sense of beau-
ty (Seghers 2015). Referring to the evolution of
adaptive mechanisms in the context of habitat se-
lection, theory and research in the field of evolutio-
nary aesthetics can explain why certain objects and/
or landscapes cause certain neural rewards and, the-
refore, a positive aesthetic appreciation. Aesthetic
reactions are also a guide to human behaviour, in-
cluding everyday behaviour with many practical con-
sequences, such as organizing work and living spa-
ces, but also in the domains such as health, design,
marketing, education, environmental protection and
the like, as previously discussed.
Every person has aesthetic preferences, but they are
not exclusively random or individual. There are uni-
versal and biophilic preferences for open grasslands,
trees, fresh water or symmetrical structures, as well
as antipathy or disgust for spiders and snakes, which
is called biophobia (Orr 1993; Ulrich 1993). All this
points to the phylogenetic basis of aesthetic reason-
ing that existed before the evolution of human cul-
ture and to the validity of evolutionary explana-
tions. In this sense, aesthetic reactions are not a tri-
vial aspect of ‘the human makeup’, nor is aesthetics
a ‘whim’ that people engage in at leisure (Kaplan
1987.26). Simply put, innate aesthetic biases helped
individuals to behave and act in an adaptive way
(Swonke 2000.260).
We have seen that numerous studies point to the
universality of landscape preferences, but it should
not be forgotten that these preferences can never-
theless be modified by cultural influences and ex-
periences, so it is not surprising that there are diffe-
rences in preferences either across (sub)cultures or
social groups (Tveit 2009). Neither biological nor
cultural reductionism is adequate, that is, biological
or social factors alone are not sufficient to explain
the individual aesthetic experience. It is not disput-
ed that landscapes are biologically important to hu-
mans and evoke certain (aesthetic) emotions, but
they are also sometimes important for one’s iden-
tity and well-being, namely, for transforming mere
habitable space into a true living place (Hunziker et
al. 2007).
The bottom line is that it is
not enough to take care sole-
ly of survival, as well-being
must also be taken into ac-
count. Still, it is critical to
note and discern the so-called
primitive preferences which
are built on human attach-
ment to nature, while in addi-
tion to the importance of (bio-
philic) design for the body,
the senses must not be ignor-
ed. Therefore, an adequate
and desirable environment is
one that takes into accountFig. 2. Charles O’Rear, “Bliss”, 1996. Landscape photography.
Marko {korić, Aleksej Ki[juhas
504
the physiological, mental and social aspects, and
that would be the evolutionary environments of
the species. They meet not only survival needs, but
also one’s well-being needs, which relate to the qua-
lity and psychological aspects of human life (Boy-
den 1971; Hildebrand 1999).
This paper was written as a part of project no. 179037
which is financed by the Ministry of Education, Sci-
ence and Technological Development of Republic of
Serbia.
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