People and Nature. 2022;00:1–18.
Received: 21 December 2021
Accepted: 29 March 2022
DOI: 10.1002/pa n3.10 330
Species diversity enhances perceptions of urban coastlines at
Tom P. Fairchild | Jasmine Weedon | John N. Griffin
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2022 The Authors. People and Nature published by John Wiley & Sons Ltd on behalf of British Ecological Society.
Faculty of Science and Engineering,
Swansea University, Swansea, UK
Tom P. Fairchild
European Regional Development Fund,
Grant/Award Number: WEFO- 80939
Handling Editor: Martin Dallimer
1. Biodiversity is increasingly understood as an important mediator of human
aesthetic appreciation of scenes and landscapes, with implications for cultural
services and well- being. However, the generality of biodiversity effects across
affective emotions, scales and habitats remains unclear.
2. Urban coastal intertidal habitats on seawalls and other artificial structures are
expanding worldwide. Despite growing calls to prioritise biodiversity in urban
coastal planning and management, the potential co- benefits determined by
people's responses to biodiversity in these novel intertidal communities are
3. We investigated, using image- based questionnaires, how several facets of bio-
diversity influence how people perceive urban coastal structures at both land-
scape and close- up scales.
4. Species richness strongly enhanced people's ratings of images for aesthetic ap-
peal, interest and calming potential at both scales, but was more pronounced at
the close- up scale. Species evenness also increased ratings at the close- up scale,
while functional diversity (Rao's Q) was associated with a decline in aesthetic
appeal and interest at the close- up scale, indicating that people can disfavour
scenes dominated by species with contrasting traits.
5. Analysis of free- text assessments showed that people strongly and positively
valued scenes that were perceived to be ‘diverse’, a response that was much
more common when viewing scenes with high species richness. The underlying
structure type also clearly affected appraisals, with more obviously engineered
structures being perceived to be less natural and thus less desirable.
6. Our results show that biodiversity's effects on aesthetic appreciation extend
to multiple affective emotions and to unfamiliar urban intertidal habitats, sug-
gesting that managing these environments for biodiversity may simultaneously
support aesthetic, educational and well- being benefits. Nevertheless, the sen-
sitivity of responses to the facet of biodiversity and viewing scale in our results
underlines the context dependency and complexity of people's perceptions of
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FAIRCHILD et al.
1 | INTRODUC TION
Humans are driving significant changes in biodiversity across eco-
logical communities globally (Chaudhary et al., 2021; Dornelas
et al., 20 14; Newbold et al., 2015), and understanding how bio-
diversity change influences ecosystem services remains a press-
ing ongoing research challenge (Balvanera et al., 2016; Gamfeldt
et al., 2013; Isbell et al., 2017). Almost three decades of research
have demonstrated that biodiversity of ecological communities
profoundly influences provisioning and regulating services such as
wild food production and carbon storage (Cardinale et al., 2012;
MEA, 2005), but growing evidence also suggests that diverse nat-
ural systems are crucial in providing valuable aesthetic and well-
being benefits (Bell et al., 2015; Ulrich, 1983; White et al., 2013).
Increasingly, researchers are beginning to investigate how biodi-
versity simultaneously influences the flow of cultural services such
as aesthetic inspiration, opportunities for tourism and recreation,
and education (Chan et al., 2012; Hoyle et al., 2017; Lindemann-
Matthies et al., 2010).
There is emerging evidence that biodiversity influences how
people perceive and respond to nature. A long history of research
in environmental psychology shows that exposure to nature can
elicit positive aspects of perception such as aesthetic pleasure
(Petrova et al., 2015; Ulrich, 1983), interest (Fairchild et al., 2018)
and a sense of calm (Dallimer et al., 2012; Fuller et al., 2007; Houlden
et al., 2018), which are key in driving inspiration, shaping positive
attitudes and enhancing psychological restoration and well- being
(Silvia, 2012; Thornhill, 2003; Ulrich, 1983). Consequently, land-
scapes that are aesthetically appealing, subjectively interesting or
providing a sense of calm are important motivators for humans to
visit and interact with natural ecosystems, while also providing sat-
isfying and engaging nature experiences (Kim et al., 2015; Pasanen
et al., 2018). Yet, rather than simply exposure to nature, it is likely
that more biodiverse and naturalistic scenes or landscapes provide
greater aesthetic appeal (Gunnarsson et al., 2017; Hoyle et al., 2017;
Palliwoda et al., 2017), and enhance well- being benefits by providing
calming spaces for relaxation and rejuvenation (Fisher et al., 2021;
Hoyle et al., 2017; Wood, Harsant, et al., 2018; Wood, Jones,
et al., 2018). However, previous studies have often taken a broad
comparative or correlational approach, with more diverse settlings
covarying with other factors, such as organism abundance, habitat
type and landscape setting, leaving the specific role of biodiversity—
and its multiple facets— poorly understood.
Indeed, it is important to consider that multiple facets of bio-
diversity may have different influences on aesthetic appeal and
other affective emotions. Most simply, increasing richness (num-
ber) of species in a community can enhance visual complexity
(Forsythe et al., 2011; Kaplan et al., 1972; Petrova et al., 2015) and
the chances that species with favourable characteristics are in-
cluded (Ulrich, 1983), intersecting strongly with empirical findings
from the psychology of aesthetics where complexity and desirable
traits can enhance aesthetic appeal (de Pinho et al., 2014; Palmer
et al., 2013; Petrova et al., 2015). Species- rich communities are also
generally perceived to be more diverse, which may lead to positive
aesthetic appraisals of communities through intrinsic existence val-
ues (Dallimer et al., 2012; Fisher et al., 2021; Schebella et al., 2019),
particularly as public awareness of the importance of biodiversity in
maintaining healthy ecosystems has increased over the past decade
(Hynes et al., 2021; Shwartz et al., 2014; Wu et al., 2020). Species
richness is, however, a coarse measure of community structure; vi-
sual complexity and diversity may be more discernible where spe-
cies abundances have higher evenness (Graves et al., 2017; Tribot
et al., 2019), or where species have more distinctive traits (char-
acteristics; e.g. colour or shape) and therefore encompass greater
functional diversity (variety of different characteristics; Fairchild
et al., 2018; Tribot et al., 2016, 2019).
Moreover, different affective emotions can be expected to vary
in responses to biodiversity. For example, increasing visual complex-
ity (Fry et al., 2009; Silvia, 2006; Sun & Firestone, 2021) and the
inclusion of a variety of species with novel, unusual traits (e.g. high
functional diversity) may elicit fascination or interest in a commu-
nity by providing unfamiliar stimuli (Fairchild et al., 2018; Kashdan &
Silvia, 2009), ultimately triggering fascination (Ballantyne et al., 2007;
Krapp, 1999; Renninger et al., 2014) and facilitating learning expe-
riences (Silvia, 2001, 2006, 2008). Conversely, restorative bene-
fits may be enhanced in communities with high species evenness
which may provide more visually coherent scenes (Fry et al., 2009;
Kang & Kim, 2019; Zhang et al., 2013), or alternatively may be in-
creased through increasing perceived health, ecological quality or
naturalness of habitats (Fisher et al., 2021; Marselle et al., 2016;
Wood, Harsant, et al., 2018; Wood, Jones, et al., 2018), which
can be driven through higher species diversity (Hoyle et al., 2019;
Purcell & Lamb, 1998; Winter, 2012). Collectively, these properties
can provide calming spaces which in turn promote self- reflection
and mental restoration (Kaplan, 2001; Pálsdóttir et al., 2018), re-
duce stress (de Kort et al., 20 06; Pálsdóttir et al., 2018; Van Den
Berg & Custers, 2011) and ultimately enhance mental and phys-
ical well- being (Garrett et al., 2019; Houlden et al., 2018; Koss &
King sley, 2010; Schwind et al., 2 017).
People also interact with natural environments at multiple scales,
from viewing a landscape while walking or travelling, to closely ex-
ploring an environment to find organisms of interest. Although
yet to be addressed, biodiversity effects on different aspects of
human– nature perceptions are likely to be highly scale dependent.
For instance, the role of abundant or conspicuous species and their
differences may be most important when viewed from afar, with
aesthetics, biodiversity, coastal structures, cultural ecosystem services, interest, seawalls,
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FAIRCHILD et al.
the total richness of organisms more discernible at close- up viewing
The role of biodiverse, naturalistic spaces for providing aes-
thetic and well- being benefits is particularly important in urbanised
areas, where opportunities to encounter nature are often limited.
Here, important nature interaction experiences for residents centre
around urban green and blue spaces, such as gardens, parks, lakes
and, increasingly, coastal spaces (Chiesura, 2004; Mills et al., 2019;
Palliwoda et al., 2017). But coastal spaces in particular, which are
increasingly becoming recognised for their potential well- being ben-
efits (Bell et al., 2015; Bell et al., 2020; Wheeler et al., 2012), are
undergoing rapid and sustained change as coastal populations con-
tinue to grow (Barragán & de Andrés, 2015; Neumann et al., 2015),
and the subsequent proliferation of urban coastal structures places
them in greater intersection with local communities and tourists
(Evans et al., 2019). This growth of fringing coastal infrastructure
and hard engineering can substantially alter the visual properties
of coastal vistas at both the landscape scale and more intimate
close- up scales (Burak et al., 2004; Morgan, 1999b) through changes
in shoreline structural complexity (Lawrence et al., 2021), with these
simplified environments typically hosting depauperate ecological
communities (Bulleri & Chapman, 2004; McKinney, 2006; Momota
& Hosokawa, 2021). This has led to increasing calls to develop
ecologically sensitive designs or management scenarios to retro-
fit coastal infrastructure, embracing varied naturalistic features to
provide suitable habitat to support natural, biodiverse ecological
communities (Evans et al., 2019; Firth, Schofield, et al., 2014; Firth
et al., 2016). Indeed, features, such as rockpools (Evans et al., 2016;
Hall et al., 2019) and surface roughness (Loke & Todd, 2016; Matias
et al., 2 010; Zawada et al., 2010) associated with older and eroded
structures— or via ecological retrofitting— may markedly increase
biodiversity along urban coastlines. In turn, the enhanced biodi-
versity may provide greater opportunities to explore and interact
with organisms at local scales (Martens, 2016; Morgan, 1999a),
together with creating more natural coastal landscape aesthetics
(Cordell et al., 2017 ). But despite the potential biodiversity benefits
provided by ecologically sensitive coastal management, there is a
lack of knowledge as to how local communities and visitors— whose
buy- in may be crucial in determining the successful implementation
of conservation or enhancement schemes— might perceive more
biodiverse urban coastal structures.
In urban coastal environments, the role of biodiversity in provid-
ing aesthetic and well- being experiences remains untested. Previous
anecdotal or implicit evidence from aesthetic appraisal studies on
artificial coastal structures— centred on how natural or congruent
they are with natural shorelines— are often contradictory (e.g. Bell
et al., 2020; Koutrakis et al., 2011; Morgan, 1999b), suggesting that
traditional positive diversity perceptions may not apply universally
across to coastal urban environments where clean architectural
lines may be sometimes preferred (Bosman et al., 2016; Koutrakis
et al., 2011). For other aspects of perception, such as interest and the
ability to provide calming spaces, experimental evidence from coastal
systems on biodiversity– benefit relationships is notably lacking, yet
is becoming particularly pressing as we begin to understand the
importance of natural blue systems for multiple cultural and well-
being functions (Bell et al., 2020; Evans et al., 2019). Furthermore,
there is limited evidence to suggest that many dominant intertidal
species, notably seaweeds, can be negatively perceived by resi-
dents and visitors at coastal locations, providing hazards to access
and egress, being associated with unfavourable aesthetics, smells or
touch sensations (Limburg et al., 2010; MacLeod et al., 2002; Merkel
et al., 2021). These potential negative perceptions of ecologically
important, characteristic species may undermine any positive bio-
diversity effects— coupled with an overall lack of understanding of
coastal biodiversity– perceptions relationships— leaving considerable
uncertainty around the aesthetics and well- being benefits created
by diversity in coastal ecosystems.
Here, we address the role of biodiversity in mediating human
perceptions of artificial coastal structures. We assessed the role
that species and functional diversity play in driving aesthetic appeal,
human interest and creating relaxing or rejuvenating environments
using an image- based questionnaire. We used a combination of
image choice, rating scales and free- text motivation tools to under-
stand the effect of biodiversity on different facets of well- being, and
to explore the underlying motivations. Furthermore, we examine
whether the scale which people view or interact with these environ-
ments— at both intimate, close- up scales associated with exploration
of nature (such as rockpooling), and at whole structure scales where
passive viewing typically occurs— may alter biodiversity– public per-
2 | METHODS
2.1 | Participant selection
Participants between the ages of 18 and 60+ were recruited
through online social media channels and promotions which linked
to the survey between 15th May and 28th September 2020. The
survey was targeted at UK and Ireland residents as part of the
Ireland- Wales ECOSTRUCTURE project, and entries submitted
from participants outside of the geographical range were disre-
garded to reduce the influence of regional variation in attitudes.
Pa rticip ant s were in for med tha t th e pu r p ose of the experim ent wa s
to ‘understanding attitudes towards coastal defence structures
and the animals and seaweed that grow on them’, but the specific
aims were not explicitly stated to prevent bias. Participants were
asked whether they consente d to being included in the study using
question- routed checkboxes (‘I consent to be part of the study’,
or ‘I DO NOT consent to be part of the study’— which routed to
the exit page) before they progressed, and informed that they
were free to withdraw, without penalty, at any time using a with-
draw (‘Exit’) button at the top of the navigation page. The survey
took approximately 15 min to complete. This study was granted
ethical approval by the Swansea University Ethics Committee
(SU- Ethics- Staff- 260,520/246).
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FAIRCHILD et al.
2.2 | Experimental design
For our study, we selected three distinctive yet complimentary
aspects of perception: aesthetic appeal, interest, and providing a
sense of calm. These aspects of perception have been identified as
important determinants of the overall motivation to visit or interact
with— and subsequent visitor satisfaction towards— natural environ-
ments (Dorwart et al., 2009; Kim et al., 2015; Pasanen et al., 2018).
To elucidate how components of biodiversity influence these dif-
ferent indicators of cultural services (aesthetic appeal, interest and
calming potential), we created an online study on Survey Monkey
(see example questionnaire section in the supplementary materials)
using images composited from photos of three different types of ar-
tificial coastal structure types and 10 species of intertidal organisms
which are common throughout the north- east Atlantic (Hawkins
et al., 2019; Table S1). The study was conducted at two focal scales:
a whole- structure scale (~c. 30 m × 5 m) and a close- up scale (~c.
1 × 1 m). Within the images, we directly manipulated species richness
(6 levels; 0, 1, 2, 4, 6, 8 species) and indirectly manipulated species
evenness (Pilou's evenness) and functional diversity (here meas-
ured as Rao's Q— B o t t a - D u k á t , 2005) through varying the commu-
nity compositions within each richness level. Species richness was
hypothesised to increase appeal, interest and sense of calm as we
expected that naturalness and complexity, which can be enhanced
by having a greater number of species, may be important in driv-
ing these perceptions. Functional diversity was included as we hy-
pothesised that greater trait differences would lead to more visually
obvious differences between species, and thus increase the ability
of participants to discriminate between species. The traits used to
generate the functional diversity measure were selected to broadly
capture differences in visual properties— such as size or colour— and
are presented in Table S2.
To assess aesthetic appeal, we used multiple different question
types to understand preference order and self- reported appeal
(Table S3). First, each participant was presented with 15 paired
image preference questions with two randomised images, side- by-
side, and asked to select which image they found most appealing (or
if they found each image to be equally as appealing). Paired image
preference approaches have been found to capture preference re-
liably and repeatably (Clark et al., 2018; Sankaran et al., 2021) and
are less susceptible to directionality biases or central tendencies
than other rating scales. As aesthetic appeal was not expected to
be the only cultural service indicator affected by diversity, we also
asked participants Likert- type questions on additional self- reported
perceptual components of how interesting (‘I find the structure in-
teresting to look at’) and calming (‘I find the structure relaxing and
calming to look at’) participants found each of the image, which have
been, respectively, seen to capture interest and the motivation to
learn (Fairchild et al., 2018; Moss & Esson, 2010; Turpie, 2003), as
well as providing relaxation and stress recovery (Christopher, 2019;
de Kort et al., 2006; Moran, 2019; Van Den Berg & Custers, 2 011).
Additionally, a Likert- type question on aesthetic appeal (‘I find the
structure appealing to look at’) to facilitate direct comparisons
between the different components and allow for the testing of de-
mographic effects on appeal. Participants were provided with 10
single images and rated aesthetic appeal, interest and calming using
a 5- point scale, including a central ‘neutral’ option (e.g. Strongly
Agree (5), Agree (4), Neutral (3), Disagree (2), Strongly Disagree (1);
Table S3). Due to layout restrictions in the Survey Monkey platform,
these three Likert- type questions were presented as a random order
question matrix. To assess any influence of acquiescence bias, we
validated results against a subset of participants with inversely
worded statements and ranked ELO results, finding no evidence of
systematic bias from positive question wording (see ‘Assessing ac-
quiescence and directionality biases from Likert- type scale data’ in
the Supplementary Materials for more information).
We also asked demographic questions (age, gender, educa-
tional/professional background, ocean connectedness (after Nuojua
et al., 2021, in review and Schultz, 2002) to understand whether
demographic differences modified relationships. Participants were
given the opportunity to not disclose their answer (‘prefer not to
say’) for each question. Further details are shown in Table S4.
Finally, to understand the motivations, participants were asked
to rate aesthetics using a Likert- type object and provide free- text
comments on what they liked and disliked about three randomised
images (‘Thinking about this structure, what motivated you to find
the structure unappealing or appealing? Please write, In your own
words, in the box below’). These questions were presented after the
demographic questions and were optional to complete.
2.3 | Image composition
We created composite images using photographs of different urban
coastal structure types: stepped concrete walls, rip- rap (or ‘rock
armour’) and an old heritage stone seawall (circa 1800s), and con-
tained 10 common and ubiquitous north- east Atlantic intertidal spe-
cies, comprising of six seaweed species, and one species each from
the groups of barnacles, anemones, limpets and mussels (Hawkins
et al., 2019; Table S1). While structure and species selections were
not exhaustive and may not fully represent the diversity present
across the country, they were chosen to represent the continuum
of complexity in UK coastal artificial habitats and communities.
Composites were created for both whole structure (c. 30 m × 5 m)
and close- up (c. 1 × 1 m) focal scales and contained either 0, 1, 2, 4,
6 or 8 species. At the structure scale, we simulated randomly se-
lected species combinations, across three different structure types,
giving a total of 120 unique images. At the close- up focal scale, the
number of structure types was reduced to two, as the riprap and old
wall sections looked too similar, giving 85 unique images. Images of
artificial structures were carefully manipulated to remove any exist-
ing seaweed or animal cover. Multiple photographs for each of the
10 intertidal species were taken across the south Wales coastline
(UK) on different substrate types, and for different orientations and
presentations. Both the structures and organisms were then digitally
extracted from the images using the scissor lasso tool in GIMP 2.9.1.,
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FAIRCHILD et al.
appropriately scaled, and layered to create images which varied in
their community composition and diversity (Figure 1). Proportions
and zonation of species were selected to realistically represent
naturally occurring communities. Care was taken to ensure no large
systematic differences in community coverage were present across
the different diversity treatments or structure types, and the per-
centage of occupancy of the structure was calculated to control for
residual variation. Structures were then resized to ensure the same
total coverage in the images. Differences in the visual properties
of wider viewsheds can affect overall aesthetic appraisal of scenes
(Morgan, 1999b; Ulrich, 1983), so here, to focus on the role of dif-
ferent facets of biodiversity and to avoid biases due to differences
in the landscape setting in which the structures were located, com-
posite structures were displayed over a separate coastal background
image which was held constant across structure types.
2.4 | Analysis
2.4.1 | Structural and image properties
The community structure within each image was quantified by ex-
amining percentage cover on the structure for each species in each
image using the Vidana image analysis tool (Hedley, 2013). From
the species- level data, we also calculated substrate occupancy by
summing visual cover to calculate the proportion of the structure
that was occupied by seaweeds or animals. Additionally, we quan-
tified the perceptual colourfulness of the image— which has previ-
ously been observed to influence aesthetic appraisals (Fairchild
et al., 2018; Tribot et al., 2 016)— using a colourfulness algorithm de-
veloped by Hasler and Suesstrunk (2003) within the Matlab R2017A
2.4.2 | Statistical analysis and visualisation
Analyses of data were performed in r statistical computing soft-
ware 3.6.3 (R Core Team, 2013). To understand how diversity
components and structural properties affected aesthetic appeal,
we created aesthetic scores from image pair win/loss/draw data
using the ELO algorithm package (Elo, 1978; Heinzen, 2017; fol-
lowing Tribot et al., 2016) to rank images. To verify that image
choices were not random, we compared the distribution of the
observed ELO aesthetic scores to randomly recompiled win/loss
dummy data using a Kolmogorov– Smirnov test. Species richness
and Pielou's evenness were calculated using the veg an package in
r (Oksanen et al., 2017) and functional diversity (Rao's Q, Botta-
Dukát, 2005) was calculated using the fd package (Laliberté &
FIGURE 1 Examples of images of structures (a– c) and close- ups (d– f) used in the survey. Images a– c include variation in the underlying
structures, from stepped walls (a) to rip- rap walls (b) and heritage stone walls (c). Images d– e show example close- up images with different
underlying structures of rip- rap walls (d and e) and stepped walls (f)
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FAIRCHILD et al.
Legendre, 2010). Rao's Q was selected as the sole measure of
functional diversity in the study as it has been found to perform
well for describing average community trait dissimilarity (Botta-
Dukát, 2005), which we expected would be a crucial factor for
people to discriminate between species. Rao's Q can also be calcu-
lated for communities containing ≥2 functionally distinctive spe-
cies, and thus could be calculated for the majority of communities
contained in our survey.
To analyse the roles of biodiversity facets (species richness,
species evenness, functional diversity), structure type and over-
all image properties (colourfulness, substrate occupancy), we
used two complimentary approaches. For the paired image pref-
erence data, we used log- linear regression mixed effects models
for analysing ELO aesthetic score relationships, with the indi-
vidual respondent as a random factor to account for different
baseline appreciation or interest in coastal communities, and in-
teractions which were expected to be important (e.g. the role of
species diversity across different coastal structure types) were
explicitly included within models. Cumulative Link Mixed Models
(CLMM) with a logit- link function— via the o rdi nal package in r
(Christensen, 2018)— were used for Likert aesthetic, interest
and calming potential relationships, again with respondent as a
random factor, and interactions included between key, interact-
ing predictors. The individual- based modelling approaches with
CLMMs also included participant demographics as predictors
to understand the generality or specificality of relationships.
Although every care was taken to capture representative survey
populations, we also statistically controlled for demographic ef-
fects by including demographic data in our CLMM models, but
some residual effects or uncaptured demographic properties,
may still slightly influence ratings. Resulting model partial ef-
fects, after controlling for other variables, were then visualised
using the plotting packages visreg (Breheny & Burchett, 2016)
and ggplot2 (Wickham, 2016). We also examined whether spe-
cies identity effects altered aesthetic appraisals using log- linear
mixed effects models (using the ‘lmer’ function in lme4 r pa ckage,
Bates et al., 2015) at both focal scales— using abundances for
the 10 species— along with the covariates of species richness and
structure type as a random factor to control for diversity and
2.4.3 | Textual analysis
As well as the numerical collection of data, optional free- text ques-
tions were included to explore the motivations of participants to
find structures appealing or unappealing. As free- text responses
can feature different words or phrases by participants to describe
the same thing, we identified motivational themes, that is, words
corresponding to broad themes within the free- text answers. For
instance, participants interchangeably used the words or phrases
‘unnatural’, ‘not natural’ and ‘artificial’ to describe the aesthetics
of the structures, which were subsequently collapsed to the com-
mon theme of ‘Artificial’, as the use of the words has the same broad
meaning. These themes were assessed by manually reading through
responses and categorising individual observations by participants,
adding new themes where observations did not fit into an existing
category. These identified themes were then visualised using a word
cloud, with the size of text scaled from the natural logarithm of the
number of responses in each category using Adobe Illustrator (CC
3 | RESULTS
3.1 | Ecological communities
A to t al of 937 people fro m the UK and Ire lan d complet ed the im age
preference sections of the sur vey (for ELO analysis), and 754 com-
pleted the image preference including demographics (CLMM anal-
ysis). Participants responded strongly to species richness at both
whole structure and close- up viewing scales, finding more spe-
cies rich and even communities on the presented seawalls to be
more app ealing (Figure 2a,b), but also surprising ly found that mo re
functionally diverse communities were less appealing (Figure 2c,f,
Tables 1 and 2, Tables S5 and S6), interesting (Tables S7 and S8),
and calming (Tables S9 and S10). The strength of these relation-
ships was stronger in close- up images than at the whole structure
scale. We found that the colourfulness of scenes positively influ-
enced aesthetic appeal, and calming potential at close- up viewing
scale s (Tables S6 and S10) but had no apparent effect at the whole
structure scale (Tables S5 and S9). We also found limited evidence
suggesting that human preferences were driven by the compo-
sition of species, finding only a weak negative effect of mussels
(Mytilus) and anemones (Actinia) at the structure- scale (Table S11),
and weak positive effect of a red alga (Palmaria) at closer viewing
distances (Table S12). As well as the effects of the ecological com-
munity there were strong preferences for the underlying structure
type. The less regular old stone wall and riprap structures were
far more appealing than the more regular concrete stepped wall
(Table 1; Figure S1).
Participants also generally found diverse structures more in-
teresting and calming than those with fewer, and less even, species
(Figure 3; Table 3). Interest was particularly strongly related to spe-
cies richness, and positive effects on interest accumulated faster
than either aesthetic appeal or sense of calm. Structure type had
a strong influence, with the more obviously artificial stepped con-
crete walls being found as less appealing, interesting and calming
compared to the old wall and riprap structures (Table 3). As with
aesthetic appeal, the positive relationships between diversity and
interest/calming were more apparent at closer viewing distances
than at the whole structure scale (Figure 3d,e,f). At the close- up
scale, we also observed negative effects of functional diversity for
interest (Table S8).
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FAIRCHILD et al.
3.2 | Participant motivations
The previous analysis showed that participants generally preferred
biologically diverse communities with less regular underlying struc-
tures. We next conducted an analysis of free- text data to better
understand what drove people's preferences for images of the pre-
sented artificial structures. In all, 500 people answered optional
free- text questions, and generated 1333 valid text responses. We
found 24 positive and 22 negative motivational themes for finding
structures aesthetically appealing or unappealing. These included
the positive attributes of ‘natural’, ‘diverse’, ‘colourful’, ‘exploration’
and ‘interesting structure’, as well as negative aspects such as ‘in-
accessibility’, seaweed- related associations (‘slimy’, ‘dirty’ and the
presence of ‘green’ algae), perceptually ‘hard’ structures, as well as
the lack of diversity in ‘depauperate’ (low diversity) communities.
Of these motivations, naturalness and diversity were the two most
mentioned positive and negative drivers of appeal by far (Figure 4;
Table S13), and were strongly related to reported aesthetic appeal
FIGURE 2 Species diversity, but not functional diversity, effects drive aesthetic preferences on artificial structures, and strengthen
with decreasing viewing scale. The effect of species richness (left), evenness (Centre) and functional diversity (right) on aesthetic appeal
scores at both the structure (a– c) and close- up scales (d– f). Centre lines on panel a and d represent the median, inner boxes represent 25th–
75th percentile and whiskers represent the maximum and minimum values. Points and lines on panels b, c, e and f represent model partial
residuals and model fits, respectively. Asterisks represent significance at α = 0.05 (*), 0.01 (**) and <0.001 (***), and ‘ns’ represents non-
Estimate SE p- value
Intercept 6.350 0.563 <0.001***
Species richness 0.068 0.013 <0.001***
Species evenness 0.036 0.022 0.102
Functional diversity (Rao Q)−0.305 0.170 0.076
rip- rap structure −0.037 0.044 0.393
Stepped concrete structure −0.084 0.033 0.012*
Occupancy 0.029 0.082 0.728
Species richness: rip- rap structure 0.014 0.017 0.385
Species richness: stepped concrete
−0.010 0.016 0.547
TAB LE 1 Species richness and
structure type influence aesthetic appeal
at the whole structure scale. Regression
model summary table examining the
role of biodiversity, structure and image
properties in driving aesthetic appeal
scores (ELO). Asterisks for p- values
represent significant results at α = <0.05
(*) and <0.001 (***)
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FAIRCHILD et al.
scores (Figure 5a,b). These motivations appeared to largely explain
people's preferences of structure type and species richness. We
found strong effects of structure type on mentions of naturalness
vs. artificialness, with greater mentions of naturalness associated
with the old wall and riprap shores, particularly as these shore types
were perceived as appealing or strongly appealing. Species rich-
ness was only very weakly associated with mentions of naturalness
(Figure 5c). On the other hand, positive mentions of diversity were
associated with positive appeal rating irrespective of structure type
(Figure 5b), while species richness was clearly associated with in-
creased mentions of diversity. Thus, the influence of structure type
appeared to be largely determined by a perception of naturalness,
while the influence of species richness was determined by a positive
perception of diversity.
3.3 | Demographic effects
Of the 937 people that answered the image questions in the sur-
vey, 754 people completed the demographics section. The analysis
of demographic effects is presented above for the whole structure
scale (Table 3), and full results table are provided in Tables S5, S7,
S9 and S6, S8, S10 for structure and close up scales, respectively.
Participants identifying as ‘female’ were overrepresented in the
survey (71%; Table 4), but other demographic groups were well
represented. While every care was taken to recruit a wide cross-
section of the public, people working in biological, environmental
or maritime roles were proportionally overrepresented compared
to the UK and Irish general population (35% of sample). Within
our sample population, we observed that demographic differences
Estimate SE p- value
Intercept 7.1 0 9 0.121 <0.001***
Species richness 0.068 0.019 <0.001***
Species evenness 0.079 0.034 0.023*
Functional diversity (Rao Q)−1.113 0.385 0.076
Stepped concrete structure −0.154 0.034 0.393
Occupancy 0.213 0 .157 0.728
Species richness: stepped concrete
0.047 0.020 0.385
TAB LE 2 Different facets of
biodiversity and structure type influence
aesthetic appeal at close- up viewing
scales. Regression model summary
table examining the role of biodiversity,
structure and image properties in driving
aesthetic appeal scores. Asterisks for p-
values represent significant results at α =
<0.05 (*) and <0.001 (***)
FIGURE 3 Species richness positively influences aesthetic appeal (left), interest (Centre) and sense of calm (right) at the structure scale
(a– c), and increases in strength at close- up (d– f) viewing scales. The percentage of respondents from 5- Likert agreement categories (strongly
disagree– strongly agree)
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FAIRCHILD et al.
between participants led to slightly different participant aesthetic
appeal (Tables S5, S7 and S9). Participant age had a slightly negative
effect on aesthetic, interest and sense of calm appraisals, with older
participants more likely to award lower scores, whereas increases
in self- reported ‘ocean connectedness’ led to participants awarding
slightly higher aesthetic appeal scores, but did not influence interest
and calming scores. For aesthetic appeal, participants that reported
high levels of ocean connectedness also responded more strongly
and positively to species richness of communities than those repor t-
ing lower ocean connectedness. However, participants working in
environmental or biology jobs were slightly more likely to award
lower aesthetic appeal and sense of calm scores than those who
did not work in an environmental industry. At the close- up scale,
however, we found demographic effects did not influence any of the
perception metrics (Tables S6, S8 and S10), and gender identity had
no effect at either scale.
4 | DISCUSSION
Despite growing interest in how biodiversity affects cultural ser-
vices, there have been few experimental tests of the role biodi-
versity's facets play in determining people's affective emotional
responses to ecological communities. Furthermore, whether
biodiversity elicits positive perceptions in more peripheral or
unfamiliar habitats remains uncertain. Our experiment test, set
across globally proliferating coastal urban environments, shows
that biodiversity affects multiple affective emotions, but that
these influences depend on both the biodiversity facet and view-
ing scale. Specifically, species richness and evenness increased
aesthetic, interest and calming qualities, while functional diver-
sity reduced these responses, and all influences of biodiversity
were stronger at the small— close- up— scale. Alongside biodiver-
sity, less uniform and more complex structures were preferred.
Estimate SE p- value
Aesthetic appeal (n = 3765)
Species richness 0.893685 0.185244 <0.001***
Stepped concrete wall (structure) −1. 3 4 3 21 0.286337 <0.001***
Participant age −0.35316 0.12 2385 <0.001***
Works in industry −0. 24605 0.102839 0.0163*
Ocean connectedness 1.38 458 0.694684 0.0462*
Species richness: ocean connectedness −1.12187 0.435947 0.0100**
Interest (n = 3770)
Species richness 0.716235 0.195321 <0.001***
Stepped concrete wall (structure) −2.306 54 0.300249 <0.001***
Participant age −0.35187 0.138959 0.0113*
Species richness: stepped concrete wall 0.325097 0.159688 0.0417*
Sense of calm (n = 3770)
Species richness 0.490491 0.182809 0.0070**
Stepped concrete wall (structure) −1.61119 0.290972 <0.001***
Participant age −0.2915 0.134378 0.0300*
Works in industry −0.3003 0.11293 0.0078**
TABLE 3 Species richness enhances
aesthetic appeal, interest and sense
of calm provided by coastal scenes at
the structure scale, whereas artificial
stepped concrete structures reduced
benefits. Model summaries for significant
predictors of appeal, interest and
calm from the Likert CLMM models,
including demographic effects from 754
participants. Full model summary tables
can be found in Tables S5– S10). Asterisks
for p- values represent significant results
at α = <0.05 (*), <0.01 (**) and <0.001
FIGURE 4 Perceived naturalness and diversity were the most important drivers of aesthetic appeal of artificial structures. The word
clouds visualise the number of mentions of different motivations for finding a structure appealing (panel a, blue hues) or unappealing (panel
b, red hues). Letter size indicates the number of mentions. Underlying data are presented in Table S12
People and Nature
FAIRCHILD et al.
Finally, our text analysis suggests that species diversity has
positive effects by increasing perceived ‘diversity’, while irregu-
lar underlying structures illicit positive responses by appearing
more ‘natural’. Collectively, these results imply that managing
and designing urban coastal environments to encourage more
biologically diverse and naturalistic environments, particularly
with irregular structures that support more species, will serve to
enhance the flow of cultural services.
How biodiversity of coastal structures is perceived by peo-
ple was unknown and we suspected that the presence of dis-
liked or unfamiliar species might generate weaker or even
negative relationships between diversity and aesthetic appeal,
FIGURE 5 Positive mentions of the important motivators of ‘naturalness’ and ‘diversity’ increase in line with appeal, while positive
mentions of ‘diversity’ increased with the species richness of communities. The top two panels represent the proportion of respondents
mentioning (a) naturalness/artificialness or (b) diversity/lack of diversity as their motivations for finding scenes aesthetically appealing or
unappealing. The bottom two panels represent the role of species richness in driving (c) perceived naturalness and (d) perceived diversity
reported by participants
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FAIRCHILD et al.
if not interest. However, despite negative associations with sea-
weed species by participants— and previous evidence of negative
seaweed associations (Limburg et al., 2010; MacLeod et al., 2002;
Morgan, 1999b)— we found a strong positive effect of species
diversity on the aesthetic appeal of common coastal structures,
which persisted regardless of demographic background. The pres-
ence of this diversity– aesthetic relationship suggests that diver-
sity effects can override potentially negative identity effects. The
diversity– aesthetic relationship also appeared to be driven by
speciose structures being accurately perceived as more diverse—
even at the structure scale where species differences were hard
to discern— with perceived diversity found to be a key driver of
aesthetic appeal from our textual analysis. These findings are in
line with previous work in terrestrial systems (Dallimer et al., 2012;
Hoyle et al., 2017; Southon et al., 2018), where perceived diversity
and naturalness have been seen to drive appeal and engagement
(Dodds et al., 2010; Turpie, 2003).
The positive effects of richness and evenness also extended to
other affective emotions of interest and calming potential, which
can mediate educational (Kashdan & Silvia, 2009; Krapp, 1999;
Renninger et al., 2014) and well- being (Cracknell et al., 2 016; Fisher
et al., 2021; White et al., 2 017) cultural ecosystem service benefits.
Self- reported sense of calm increased similarly to aesthetic appeal in
our study despite a exhibiting a slightly lower baseline calming po-
tential than was observed for appeal. Previous studies have indicated
that restorative calming properties of scenes or landscapes may be
strongly linked to aesthetic appeal, as the same properties such as
naturalness and ecological quality are thought to be important in re-
ducing stress induced by the psycho- evolutionary appraisal of habi-
tat suitability, favouring perceptually ‘healthy’ ecosystems (Bratman
et al., 2019; Kaplan, 1987; Orians & Heerwagen, 1992). Importantly,
species diversity can substantially contribute to human appraisals
of the perceptual naturalness and ecological health of landscapes
(Barker & Fisher, 2019; Rapport et al., 1998; Tribot et al., 2018) and
as such may promote a sense of calm and relaxation in people, ulti-
mately reducing stress (de Kort et al., 2006; Fritz et al., 2 010; Van
Den Berg & Custers, 2011) and providing mental and physical well-
being benefits (Gu et al., 2015; Pálsdóttir et al., 2018).
Interest, in particular, increased strongly with species richness
and accumulated more quickly with diversity than aesthetic appeal
or sense of calm. This rapid accumulation of interest with increasing
diversity was likely driven through a combination of increasing visual
diversity, complexity and increasing probability of including novel
species— mechanisms which have been previously demonstrated to
drive interest (Kashdan & Silvia, 2009; Krapp, 1999; Silvia, 20 08)—
and an increase in participant reported desire to explore the ‘un-
seen’ diversity potential which can be offered by seaweed canopies
(Martens, 2016). These diversity– interest relationships are ex-
pected to provide strong motivations for exploration and facilitate
learning experiences (Ballantyne et al., 2007; Fairchild et al., 2018;
Simmons, 1998), ultimately providing educational opportuni-
ties (Kashdan & Silvia, 2009; Krapp, 1999; Renninger et al., 2014).
Furthermore, this strong diversity– interest relationship, and the ap-
parent decoupling of the magnitude of diversity– interest relation-
ships from calming potential and the better understood aspect of
aesthetic appeal (Cupchik & Gebotys, 1990; Kashdan & Silvia, 2009;
Turner & Silvia, 2006), suggests that diversity independently drives
multiple aspects of perception— and therefore supporting cultural
and well- being services— through different mechanisms.
How biodiversity influences the functions and values of ecosys-
tems has been a highly active area of research in ecology (Balvanera
et al., 2016; Gamfeldt & Roger, 2017; Lefcheck et al., 2015), with
theory suggesting these relationships can be strongly scale de-
pendent (Isbell et al., 2017; Pasari et al., 2013). Our work shows
that scale dependence also extends to diversity– aesthetics and
diversity– interest relationships. Indeed, we found that viewing scale
determined the impact of several facets of biodiversity on aesthetic,
and interest benefits. Previous studies have tended to focus on
TAB LE 4 Demographic background summaries for the
participants for the online study who completed the demographic
Other gender identity 3
18– 30 166
31– 45 278
46– 60 221
Frequency of coastal visits
I have never been to the coast 1
I rarely visit the coast 153
I visit the coast at least once a month 201
I visit the coast at least once a week 129
I live on the coast 270
Work in environmental related job
Yes, for a port authority or in port operations 8
Yes, in a regulatory environmental role 50
Yes, in biological sciences 214
Yes, in civil engineering 25
Ocean connectedness (see Table S3)
No connection 58
Some connection 295
Strong connection 290
‘As one’ 106
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FAIRCHILD et al.
broader, feature or landscape scale, effects of biodiversity (Dallimer
et al., 2012; Lindemann- Matthies et al., 2010; Tribot et al., 2019),
but close- up scales are also relevant because coastal structures are
frequently used intimately, providing a setting for exploration and
learning opportunities through close examination of the rocks and
ecological communities which can inhabit them (Martin et al., 2006).
Our study found a marked strengthening of species richness and
evenness effects at the close- up scale, which was likely explained
through differences between species and communities (e.g. identity,
colour, texture and shape, dominance)— and the desirable property
of ‘diversity’ underlined in the textual analysis— being more percep-
tible at close- up viewing scales.
While species diversity strongly contributed to multiple as-
pects of perception at both the landscape scale and close- up
scale, we observed more complex relationships with functional
diversity. We found no evidence that functional diversity (mea-
sured as Rao's Q) substantially affected aesthetic appraisal, in-
terest or calming potential at the whole structure level, but we
observed stronger, negative effects at close- up viewing scales.
This suggests that the inclusion of organisms with the extreme
trait differences may be undesirable at close viewing scales where
species (and trait) differences can be more readily resolved, ei-
ther through species with contrasting extreme traits being disliked
(Ulrich, 1983), or by exceeding some threshold of visual complex-
ity, which may lead to poorer aesthetic appraisals of structures
(Forsythe et al., 2011). Combined with recent findings from coral
reef fish assemblages that functional diversity metrics can have
weak, or hump- shaped, relationships with aesthetic preference
and be outweighed in importance by species richness (Tribot
et al., 2019), this result suggests that— unlike ecosystem func-
tions such as primary productivity, and related provisioning and
regulating services, which are often strongly explained by func-
tional diversity (Cadotte et al., 2011; Gross et al., 2017; Mensah
et al., 2020)— functional diversity and cultural services related to
aesthetics may be somewhat decoupled or even negatively asso-
ciated and may trade- off against different functions and services.
Overall, our results underline that different facets of biodiversity
can have different impacts on people's preferences and at differ-
ent focal scales, although more work is required to fully untangle
the causes and generality of these differences, including across
other systems and habitats.
Furthermore, the application of these scale- dependent rela-
tionships to other systems will depend on the spatial structure of
communities. In our model communities, species were well- mixed
spatially, such that close- up images contained a large proportion of
the overall species pool. However, in some systems and contexts,
systems can be patchy with different patches dominated by differ-
ent species such that small- scale (close- up) diversity can be much
lower than wider— landscape— diversity. In such cases, aesthetic ap-
preciation and even interest may be greater at the larger landscape
scale. Future work should explore the interaction between spatial
community organisation and scales of diversity (i.e. alpha, beta and
gamma) and human perceptions and well- being.
While more diverse structures were seen to be more natural, di-
verse and subsequently appealing, the structural characteristics of
underlying coastal structures were highly important for aesthetic
appraisals of appeal. More obviously engineered structures, such as
the concrete stepped walls, were far less preferred than those which
were perceived to be more natural. The less defined edges and facets
of the riprap and heritage stone walls led to greater perceived natu-
ralness and perceptions of diversity than for similar communities on
the hard- edged, regular stepped concrete wall structure, with many
participants no ting the app are nt ‘hardness’ of the st ructure as an un-
desirable trait which influenced their overall aesthetic appraisal. This
desire for more natural or ‘organic’ shapes is a well- understood aes-
thetic driver of building architecture (Coburn et al., 2019; Vartanian
et al., 2013), shifting baseline aesthetic appreciation, interest and
calming potential of structures— underlining the multiple ecological
and anthropogenic motivators for aesthetic appraisal, and the par-
ticular importance of structural properties which needs to be better
integrated into future natural aesthetics studies.
The strong relationships between species diversity and aesthet-
ics, interest and calming potential demonstrate the importance of
human– nature interactions that healthy, diverse ecosystems can
provide. This further strengthens growing arguments for the need
to better integrate cultural ecosystem services into the discourse
around the value of nature— which has tended to be neglected (Chan
et al., 2012; Daniel et al., 2012; Martin et al., 2016)— into coastal
management and planning activities. At the broad scale, our findings
suggest that managing coastal structures to promote diverse com-
munities is likely to positively contribute towards well- being com-
mitments set out in international and national goals and legislation,
such as the U.N. Sustainable Development Goals (United Nations
General Assembly, 2015; Wood, Harsant, et al., 2018; Wood, Jones,
et al., 2018) or the Wellbeing of Future Generations Act 2015
(Wales), as well as directly benefiting local communities. Given the
importance of aesthetics in driving attitudes and generating local
support for changes to— or creation of— artificial coastal structures,
we found strong evidence that images of biodiverse seawalls are
more desirable to communities than highly engineered, low diversity
depauperate structures, and found that these biodiversity benefits
were realised regardless of demographic background. However, it
is important to note that other aspects of nature experiences may
not be captured by visual senses alone (e.g. sound, smell or touch),
with growing evidence suggesting that aesthetic experiences and
well- being benefits can be further enhanced through non- visual
stimuli driven by increased biodiversity (Austen et al., 2021; Fisher
et al., 2021; Franco et al., 2017). This suggests that while the posi-
tive biodiversity– perceptions relationships presented here are likely
to be good indicators of human benefits, our value estimates from
image- based elicitation surveys associated with diverse coastal
communities are likely to be conservative (Fisher et al., 2021; Franco
et al., 2017; Schebella et al., 2019). Future studies, including con-
trolled scenes as well as in- situ surveys, are important to address this
uncertainty. Our results also suggest that increasing engagement
with coastal areas may also generate greater biodiversity— well- being
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FAIRCHILD et al.
benefits, with the feeling of connectedness to the sea enhancing
biodiversity– appeal relationships. Along with evidence that bio-
diverse communities can support a wide range of supporting and
regulating services which sustain resilient and healthy ecosystems
(Balvanera et al., 2016; Barbier et al., 2011; Cardinale et al., 2012),
our findings support calls to explicitly include ecologically sensitive
design into coastal defence projects, either through retrofitting
existing infrastructure to improve biodiversity outcomes (Evans
et al., 2016; O'Shaughnessy et al., 2020), or through including
biodiversity- enhancing features on new or replacement structures
(Firth, Thompson, et al., 2014; Perkins et al., 2015) to enhance well-
being ecosystem service provisions.
5 | CONCLUSION
In this study, we provide experimental evidence that multiple as-
pects of perception— which are important mediators of a wealth
of educational, aesthetic and well- being ecosystem services— are
generally enhanced by specific facets of biodiversity in urbanised
coastal settings. While limited evidence of biodiversity enhance-
ment of individual aesthetic and well- being benefits exists for ma-
rine systems, we show how despite negative preconceptions of
intertidal species, more diverse artificial coastal habitats can still
provide attractive, interesting and restorative spaces for people.
This simultaneous facilitation of the perceptions of interest, aes-
thetic appeal and sense of calm in this study, coupled with pre-
vious evidence of provisioning and regulating services, suggests
that biodiversity plays a crucial role in providing multi- functional
coastal urban spaces which maximise human benefits at multiple
scales of use. As such, management interventions that enhance
diversity on urban coastal structures may help offset some of
the negative ecological and cultural costs associated with coastal
urbanisation. However, the unexpected effects of functional di-
versity and evidence of scale dependence of biodiversity– benefit
relationships highlight those mechanisms driving often- neglected
cultural and well- being benefits need to be explored more fully to
maximise the impact of sustainable biodiversity management and
5.1 | Study limitations
A further discussion on the limitations of the methodologies
and study system is found in the ‘Limitations’ section in the
The authors would like to thank the Ecostructure project staff for
providing feedback on the survey design. This research was funded
as part of the Ecostructure project (part- funded by the European
Regional Development Fund (ERDF) through the Ireland- Wales
Cooperation Programme 2014- 2015).
CONFLICT OF INTEREST
The authors declare no conflict of interest.
T.P.F. and J.N.G. conceptually conceived and developed the study
and contributed equally to the writing of the manuscript; T.P.F. and
J.W. created the composite images, online survey templates and led
participant recruitment; T.P.F. led the analysis of survey results and
was supported by J.N.G. and J.W.
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Additional supporting information may be found in the online
version of the article at the publisher’s website.
How to cite this article: Fairchild, T. P., Weedon, J. & Griffin,
J. N. (2022). Species diversity enhances perceptions of urban
coastlines at multiple scales. People and Nature, 00, 1–18.