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Ecol Solut Evid. 2023;4:e12243.
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https://doi.org/10.1002/2688-8319.12243
wileyonlinelibrary.com/journal/eso3
Received:11November2 022
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Accepted :18April20 23
DOI: 10.10 02/2688 -8319.12243
RESEARCH ARTICLE
Urban wildflower meadow planting for biodiversity, climate
and society: An evaluation at King's College, Cambridge
Cicely A. M. Marshall1,2 | Matthew T. Wilkinson3 | Peter M. Hadfield4 |
Stephen M. Rogers3 | Jonathan D. Shanklin5 | Brian C. Eversham6 | Roberta Healey7 |
Olaf P. Kranse7 | Chris D. Preston8 | Steven J. Coghill2 | Karris L. McGonigle1 |
Geoffrey D. Moggridge2 | Peter G. Pilbeam9 | Ana C. Marza10 | Darinka Szigecsan3 |
Jill Mitchell1 | Marcus A. Hicks1 | Sky M. Wallis1 | Zhifan Xu1 | Francesca Toccaceli1,2 |
Calum M. McLennan2 | Sebastian Eves- van den Akker2,7
This is an op en access arti cle under the ter ms of the CreativeCommonsAttribution License, which permits use, distribution and reproduction in any medium,
provide d the original wor k is properly cited.
©2023TheAuthors .Ecological Solutions and Evidence publishe d by John Wiley & Sons Ltd on behalf of Brit ish Ecological Societ y.
1Conservation Research Institute and
Department of Plant Sciences, University
of Cambridge, C ambridge, UK
2King's College, U niversity of C ambridge,
Cambr idge, UK
3Depar tment of Zoolog y, University of
Cambridge, Cambridge, UK
4Ecology Solutions, Cokenach Estate,
Royston, UK
5Botanical Society of Britain & Ireland,
Cambr idge, UK
6The Wildlife Trust for Beds, C ambs an d
Northants, Cambridge, UK
7The Crop Science Centre, Department of
Plant Sci ences, University of C ambridge,
Cambr idge, UK
8Independent Researcher, Cambridge, UK
9Cambridgeshire Mammal Group,
Cambr idge, UK
10Downi ng Colle ge, Unive rsit y of
Cambridge, Cambridge, UK
Correspondence
CicelyA .M.Marshall
Email: cm997@cam.ac.uk
Funding information
Biotechnology and Biological Sciences
ResearchCouncil,Gra nt/AwardNumber:
BB/N021908/1, BB/R011311/1 and
BB/S006397/1; Gatsby Charitable
Foundat ion; King's College Cambridge,
University of C ambridge
Handling Editor: Harriet Downey
Abstract
1. The biodiversity and climate crises are critical challenges of this century.
Wildflower meadows in urban areas could provide important nature- based solu-
tions, addressing the biodiversity and climate crises jointly and benefitting society
in the process. King's College Cambridge (England, UK) established a wildflower
meadow over a portion of its iconic Back Lawn in 2019, replacing a fine lawn first
laid in 1772.
2. Weusedbiodiversitysurveys,WilcoxonsignedrankandANOVAmodelstocom-
pare species richness, abundance and composition of plants, spiders, bugs, bats
andnematodessuppor tedbythemeadow,andremaininglawn,over3 years.We
estimated the climate change impact of meadow vs lawn from maintenance emis-
sions, soil carbon sequestration and reflectance effect. We surveyed members
of the university to quantify the societal benefits of, and attitudes towards, in-
creased meadow planting on the collegiate university estate.
3. In spite of its sm all size (0.36 ha), the meadow su pported approx imately three
times more plant species, three times more spider and bug species and indi-
viduals, and bats were recorded three times more often over the meadow than
the remaining lawn. Terrestrial invertebrate biomass was 25 times higher in the
meadow compared with the lawn. Fourteen species with conservation designa-
tions were recorded on the meadow (six for lawn), alongside meadow specialist
species.
4. Reduced maintenance and fertilising associated with meadow reduced emis-
sionsbyanestimated1.36 MgCO2- e per hectare per year compared with lawn.
Relative reflectance increased by 25%– 34% for meadow relative to lawn. Soil car-
bon stocks did not differ between meadow and lawn.
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1 | INTRODUC TION
The climate and biodiversity crises are critical challenges of this cen-
tury. Urban areas have been cited as an impor tant part of both the
climateandbiodiversityproblem(Aronsonetal.,2014) and solution
(Beninde et al., 2015; Cadotte et al., 2021). Wildflower meadows are
often encouraged as part of offsetting schemes, nature recovery
plans and city greening initiatives (Bretzel et al., 2016; Her Majesty's
Government, 2019; Klaus, 2013). Wildflower meadows in urban
areas could provide important nature- based solutions, addressing
the climate and biodiversity crises jointly, and benefitting society in
the process (Stafford et al., 2021).
Although40%of landcoverintheUnitedKingdomisgrassland
(Carey et al., 2008), just 2% of this comprises biodiverse and carbon-
rich semi- natural grassland (Bullock, 2011; Norton et al., 2021).
Semi- natural grasslands declined by 97% between 1930 and 1970
due to ploughing, drainage and fertilisation (Fuller, 1987 ), with the
result that more conservation priority species are associated with
grasslands in the United Kingdom than with any other habitat type
(The Grasslands Trust, 2011). Wildflower meadows are semi- natural
grassland habitats, rich in native forbs, managed by taking a regular
hay crop with or without aftermath grazing (Rothero et al., 2016).
Not only do meadows have high biodiversity value, under the right
circumstances they can contribute to climate change mitigation and
adaptation, and improved human wellbeing (Nor ton et al., 2021).As
anthrop ogenic semi- natural habitat s, meadowlands are central in our
cultural and social history (Lewis- Stemple, 2015; Rackham, 2000).
The preservation and restoration of wild flower meadows is now of
strategic national interest (Her Majesty's Government, 2019).
In the United Kingdom, 84% of people live in urban areas (World
Bank, 2022).Anestimated25%oftheUK 'surbanareaisgrassland,of
which 64% is found in patches smaller than <2500 m2, and about half
of which is found in gardens (Evans et al., 2009). Most urban grassland
comprises lawn: short mown, species poor, Lolium perenne (perennial
rye grass) sport s turfs, or Agrostis- Festuca fine turfs (Hubbard, 1992).
Lawns have been the dominant form of urban grassland since the
20th century and are now a social norm, attributed with easy estab-
lishment, maintenance, recreational and aesthetic benefits (Hoyle
et al., 2017; Ignatieva et al., 2017; Norton et al., 2 019). By convert-
ing urban lawns to urban meadows, urban grasslands could present
a significant opportunity to integrate grassland conservation efforts
within human- dominated landscapes for the benefit of both people
and wildlife (Chollet et al., 2018; Norton et al., 2 019). The oppor tunity
is significant thanks to the low baseline biodiversity value of lawn, and
the extent of amenit y lawns across the UK's cities.
Althoughbothlawnsandmeadowsaresemi-naturalhabitatscre-
ated and maintained by people, they differ in the intensity of man-
agement regime, with lawns being more frequently mown, fertilised,
watered and applied with pesticides (Rorison & Hunt, 198 0). These
important differences in management between lawns and meadows
have consequences for biodiversity both above- ground and below-
ground, carbon sequestration,aesthetics and amenity value. Above-
ground, restored or experimental meadow plots have been associated
with high dicot: monocot ratios and high overall plant species rich-
ness (Chollet et al., 2018; Scotton & Rosset ti, 2021). This in turn pro-
motes significant richness and abundance of pollinating insect species
(Hutchinson et al., 2020), with positive consequences for higher tro-
phic feeding levels (Scherber et al., 2010). Below- ground, changes to
the soil microbial community of bacteria and fungi were observed in
recently established urban meadow plots (Norton et al., 2019), while
plant diversity and composition altered soil nematode communities in
manipulatedgrasslands(Viketoftetal.,2009). Meadows' potential role
in climate ch ange mitigation in cludes increas ed rates of carbon s eques-
tration compared with amenity grasslands: species- rich grasslands re-
stored from species- poor swards showed increased rates of carbon
sequestration compared with species poor swards ( Yang et al., 2019),
and increased plant species richness in manipulated meadow plots
(Norton et al., 2019). For people, biodiverse perennial meadows were
found to increase residents' perceptions of site quality in urban green-
space (Southon et al., 2017 ), and were considered the most attrac-
tive of a range of grassland planting schemes to respondents in China
(Jiang & Yuan, 2 017). While natural and semi- natural grasslands are
well studied, urban grasslands have received relatively less attention
from the restoration ecological community (Klaus, 2013).
The lawn probably first appeared in medieval times in Europe
(Ignatieva et al., 2017). L awn became more widely est ablished with the
5. Respondents thought meadows provided greater aesthetic, educational and men-
tal wellbeing services than lawns. In open responses, lawns were associated with
undesirable elitism and social exclusion (most colleges in Cambridge restrict lawn
access to senior members of college), and respondents proved overwhelmingly in
favour of meadow planting in place of lawn on the collegiate university estate.
6. This study demonstrates the substantial benefits of small urban meadows for
local biodiversity, cultural ecosystem services and climate change mitigation, sup-
plied at lower cost than maintaining conventional lawn.
KEYWORDS
biodiversity, climate change, lawn, nature recovery, nature- based solutions, restoration, urban
ecology, wildflower meadow
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development of the picturesque and gardenesque landscape architec-
tural st yles in the 18th and 19th centuries in Europe, the United St ates,
Australiaand NewZealandandhasbeenusedonaver ywidescalein
all green areas, public and private, since the 20th centur y (Ignatieva
et al., 2017). The city centre of Cambridge, in southern England (UK),
is aesthetically and culturally dominated by buildings associated with
the departments and colleges of the University of Cambridge, many of
which were established during the medieval period and stand today
more or less as they were f irst designed. King's College occupies a very
central position in the City, on King's Parade. The much- photographed
King's Chapel, Gibbs building and Back Lawn together comprise the
iconic and most often- used image of Cambridge (Figure 1, aerial view).
ThelandHenryVIacquiredforKing'sCollegethroughthe1440swas
already in the centre of Cambridge. This densely populated area ac-
commodated hostels, gardens, a convent, some common ground and
a churchyard with burial ground, the remains of which all lie under the
college buildings today (Willis & Clark, 1886).
Althou gh it is tempting to t hink of the land scape of colle giate
Cambridge as unchanging, in 1574, the college and city lands west
of the river now known as ‘the Backs’ was a simple landscape of
unimproved marshy pasture, which by 1592 had been formalised
with avenues of trees, an artificial pond, and a duck house. In place
of the King's Back Lawn were an orchard, a bowling green and a
fellows' garden, which were replaced in 1688 with a grass sward
divided by paths and avenues of trees. The Gibbs building was con-
structed from 1724. Not until 1772 did the idea to improve the land
between Gibbs and the river take shape; the tree lined avenues were
removed and the whole area was laid out as lawn to form the ‘Great
Square’ of grass, known today as the Back Lawn. Where the fashion
in landscapes and gardens had once been about the demonstration
of wealth via productivity, by the 1770s, good t aste now dictated
the opposite, that wealth be demonstrated by how much land one
could afford to keep out of productivit y, which was ver y much the
statement made by the Back Lawn (Willis & Clark, 1886).
In autumn 2019, King's College Cambridge sowed a wildflower
meadow in place of a portion of the Back Lawn. In designing the plant-
ing, the aim of the college governing body was to improve the wildlife
value and reduce greenhouse gas emissions, while providing greater
benefits to the college and wider Cambridge community. Here, we
evaluatetheperformanceoftheKing'smeadowusing3 yearsofdata,
in each of three areas: (1) wildlife value, (2) climate change mitigation
and (3) societal benefits. We consider the desirability and feasibility of
increased meadow planting across collegiate Cambridge and beyond.
2 | MATERIALS AND METHODS
2.1 | Study site
2.1.1 | Meadowestablishment
The meadow area covers about 40% of the original extent of the
King's College Back Lawn, which was first laid in 1772 (Figure 1). The
dimensionsare96 × 66 mlawn(0.63 ha)and96 × 37 mmeadow(0.36 ha).
Asoilstudycommissionedbeforesowingshowedboththetopsoil(of
30 cmdepth) and subsoilwerestronglyalkaline (pH 8.4)sandyloams.
The topso il had interme diate fert ility (20–27 mg/L ex trac table pho s-
phorus,131–167 mg/Lextractable potassium, 0. 50%–0.52% totalni-
trogen using Dumas method), while the subsoil had moderately high
fertility (35–54 mg/Lextractable phosphorus,69–129 mg/L extracta-
ble potassium, 0.24%– 0.39% total nitrogen). Thus, topsoil removal was
not necessary, and seed was sown into glyphosate treated scarified
topsoilat6 g/m2 in October 2019. Three different seeds mixes sourced
by Emorsgate were sown: the Great Lawn meadow mix, a perennial
meadow species mix intended as the long- term flora of the meadow;
a Cornfie ld Annual mix in tended to provid e first year col our; and a
Supplementary Mix composed of species with lower establishment
probability from seed, but high conservation value (Table S3).
2.1.2 | Meadowmanagement
The mead ow is managed as an Eas t Anglian hay mea dow following
traditional Lammas practices as far as possible. Hay is cut once a year
around August 1st (Lammas day) to aheight ofc. 350 mm, with one
subsequentcutat350 mminDecember,inplaceofthehistoricallight
grazing. Hand weeding was performed through the visitor seasons
to remove the occasional individual of undesirable species (mainly
Sonchus oleraceus and Cirsium vulgare). No other management or inter-
vention has b een practise d. Management of the r emaining 60% of lawn
continues as before; the lawn is a fine lawn mix with Agrostis stolonifera
and Festuca rubra dominant. It is maintained with twice- weekly cuts
from March to September, weekly cut s from October to December,
dropping to biweekly cuts in January and February. NPK fertiliser is
appliedatc.30 g/m2 in spring (8% N, 7% P, 8% K) and winter (3% N, 8%
P,8%K).Aselecti veherbicide(Prax ys)isappli edtoth eremai ninglawn
at the minimum dosage once to twice per year. Insec t pesticides are no
longer applied, and watering is avoided as far as possible. Fertiliser and
herbicide is applied in a directional fashion by ride- on vehicle during
suitable weather conditions only to minimise run- off.
2.2 | Biodiversity
2.2.1 | Plants
Botanic al surveys were carried out in July for each flowering sum-
mer (2020, 2021) and in September for the pre- sowing baseline
(2019). Five quadr ats 50 × 50 cm were place d every 15 m perpe n-
dicular to the edge in both the meadow and the lawn (KBME01-
KBME05, KBSO01- KBSO 05, Figure 1). The origin of the meadow
transe ct is 15 m from the nor thern lawn edge, an d 5 m from the
eastern lawn edge, at latitude 52. 204691°N, longitude 0.115580°E.
Theoriginofthelawntransectis15 mfromthesouthernlawnedge,
and5 mfromtheeasternlawnedge,atlatitude52.204045°N,longi-
tude0.115737°E.Abundancewasmeasuredbycountingpresencein
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each of 25 equal subdivisions of the quadrat. Mean plant height was
recorded. In addition, running checklists of all species present in the
lawn and meadow separately were collected over the course of each
year, with 2– 3 principle recording visits made each year in March,
AprilandJuly.Plantswereidentifiedassownornon-sownusingthe
stated seed mix (Table S3). Plant attribute data (distribution, scar-
city) wer es ourced from PL ANTATT (Hill et al., 2004). Designated
species follow JNCC (2022).
2.2.2 | Invertebrates
Above-ground invertebrates were sampled by sweep net (July
2020, July 2021, pre- mowing) and pitfall trap (September 2019,
2020, 2021, post- mowing) at five point s in both the meadow and
lawn (Figure 1). Sweep net transects were 20 paces each, centred
on the plant quadrat locations. Sweep net specimens were identified
to species level for all taxa in 2020, using morphospecies names as
nece ss ary,andtosp ec iesforHem ip ter a( bu gs)a ndAra nea e(spid ers)
onlyin2021. 2020datawererestric tedtoHemipteraandAraneae
for analysis. Pitfall traps were sited at the centre of the plant quadrat
locations. Pitfall trap specimens were weighed in 2021 only. Pitfall
specimens were identified to species for all taxa present in 2019
andtospeciesforHemiptera,AraneaeandOrthopteraonlyin2020
and 2021. Spider attribute data (hectad occurrence, habitat prefer-
ences) were sourced from British Spiders (2022). Hemiptera habitats
were sourced from British Bugs (2022), with hectad distribution data
from National Biodiversity Network (2022). Arthropod body size
data were compiled from NatureSpot, British Bugs, BugGuide and
Bugwoodwiki (2022); male and female maximum body lengths were
averaged. Designated species follow JNCC (2022).
2.2.3 | Bats
Bats were surveyed via two unattended ultrasonic recorders
(Wildlife Acous tics Song Meter SM4BAT FS Ultrason ic Recorder)
placed adjacent to the meadow, and the lawn (Figure 1). Recorders
were left for five or six nights each over four recording periods in
May, June, July a nd October i n 2021 only. Audio fil es were auto-
identified to species using Kaleidoscope version 5.4.6 before being
checked ma nually. All records of Barbastella barbastellus were ac-
cepted, one record of Myotis bechsteinii was assigned to Myotis
daubentonii; Plecotus austriacus records were assigned to Plecotus
auritus or Eptesicus serotinus, one Rhinolophus ferrumequinum record
was assigned to Pipistrellus pipistrellus. Myotis species are gener-
ally considered indistinguishable by audio recording only. The only
Myotis species recorded in our dataset was auto- identified as Myotis
daubentonii, which was also seen foraging at the river, and so the
identity has been retained for analysis. The total number of echo-
locations recorded over the year in each habitat is used as a proxy
for abundance (several passes by the same bat would not be distin-
guished). Designated species follow JNCC (2022).
2.2.4 | Soilnematodes
Soils were sampled contemporaneously with the pitfall traps in
September 2019, 2020 and 2021, and were co- located (Figure 1).
Approximately 7 cm width by 10 cm depth of soil was dug and
mixed. Ne matodes were ex tracted by wet ting 180–200 g of soil
on top of a paper towel with RO water. The wetted soil was lef t
overnight in a tray covered with an autoclave bag to prevent evap-
oration.Theflowthrough was collected in1 Lglassmedia bottles
(Fisherbrand), andleft to settle at a 45° anglefor24 h.The sedi-
mentwaspipettedintoa50 mLconical centrifuge tube(Corning)
using a soda lime glass pipette (Fisherbrand) and centrifuged at
300×RCF for 15 min. The pellet was transferred to a 1.5 mL mi-
crocentrifuge tube (Eppendorf) and centrifuged at 20,00 0 RCF
for1 minandsnapfrozeninliquidnitrogen.Thefrozentissuewas
lysed at 30 Hz in a tissue lyser(Qiagen) for 2 min withone 5-mm
andtwo2-mmglassbeads (Qiagen). From the samples, DNA was
extractedusingaChargeSwitch™gDNAMicroTissueKitminipro-
tocol.Usingthewell-established18S RNAprimers,NemFopt and
18Sr2bRopt ( Waeyenberge et al., 2 019) DNAwasamplified(Q5®
High-Fidelity DNA Polymerase) via PCR and cleaned using the
Monarch® PCR &DNACleanup Kit5 μg.TheamplifiedDNA was
sent to the GENEWIZ Takely Laboratory (UK) for next- generation
sequencing.
2.2.5 | Analysis
Mean richness and abundance of plants, spiders and bugs (pitfalls,
sweeps), nematodes and bats recorded in the meadow in 2021 were
compared with the pre- sowing meadow baseline (2019) and lawn
control (2021) using Wilcoxon signed- rank tests, paired for bats only
as observations are paired across nights, with bat activity each night
highly variable (Table 1; Figure 2). p values are reported at p < 0.001,
p < 0.01 and p < 0.05, with p < 0.05taken as significant, thatis, the
Bonferroni correction is not applied. In this case all tests are of a
priori hypotheses; while the Bonferroni correction reduces the
chance of type I errors, the likelihood of type II errors increases sub-
stantiallywiththisstringentcorrection.Vaguenameswereexcluded
from analyses and checklist totals, unless unambiguously assignable
to a unique species, for example, Zelotes sp. where no other Zelotes
species was recorded. Missing pitfall trap values were replaced with
the group mean for model testing.
Thestudyhasabefore-after-control-impact(BACI)design,with
samplinginitiatedbeforethemeadowwasestablished.BACIdesigns
are an effective method to evaluate perturbations when treatment
sites cannot be randomly allocated or blocked, as in the current con-
text (Conner et al., 2015). A signifi cant interac tion betwe en loca-
tion (control/impact) and year (before/after) allows any change in
response variables to be attributed unequivocally to the meadow
planting rather than temporal or spatial heterogeneit y. BACI hy-
pothesistestingwascarriedoutusingANOVA,withresponsevari-
ables transformed after visual inspection of the residuals showed
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MARSHALL et al.
this was necessar y; transformations used are reported alongside the
Results (Table 2). Contrasts were calculated post- hoc using Tukey
Honest Significant Difference; significant contrasts are presented in
Table S1.
Non- metric multidimensional scaling (NMDS) ordination was
carried out in r package vegan 2.5.5 (Oksanen et al., 2019) using a
Bray– Curtis dissimilarity matrix calculated from the plants, sweeps,
nematodes and bats datasets from the five meadow and five lawn
sample sites across all 3 years (n = 30) (Figure 3). Two axes were
specified (three for bats); convergence was reached within 20 runs
with stress <0.2for all analyses. A chi-squared test of association
between bat species and location was performed (Figure 5).Allan al-
yses were carried out in R version 3.6.0 (R Core Team, 2020) unless
otherwise stated.
2.3 | Climate change
2.3.1 | Emissionsreductions
Greenhouse gas emissions (CO2e) associated with the maintenance
regimes (mowing and fertilising) of the lawn and meadow were es-
timated using literature values (Table 3). The total area of greens-
pace across Cambridge College sites was estimated using computer
vision on high- resolution image exports from the Universit y Map
website (https://map.cam.ac.uk). The scale bar in the interactive
viewpor t (e.g. 50 pixels = 500 m) was noted for each image and
used to conver t area measurements from pixels to metres squared.
The method was checked for consistency against staff estimates
and a measurement from Google Earth. Map exports were pro-
duced at several scales to check for scaling errors. The commented
code and resources are hosted at https://github.com/DOD14/
map_area_calcu lator.
2.3.2 | Carbonsequestration
Soil organic matter (SOM) was measured as a proxy for soil car-
bon sequestration. Soils were sampled contemporaneously with
the pitfall traps in September 2019 and 2021 and were co- located
(Figure 1).7 cmwidthby10 cmdepthofsoilwasdugandmixed.For
SOM, 100 g of soil fromeach samplewasdried at 70°Cfor2 days,
homogenised and sieved(2 mm),then weighedinto threepseudor-
eplicat es of 5.00 g each p er sample loc ation. SOM was e stimated
usingthelosson ignitionmethod: samplesweresubjectedto8 hin
a muffle furnace at 450°C and reweighed once cool (Pribyl, 2010).
SOM for the meadow and lawn samples were normally distributed
and were compared using a t- test. We used a conversion factor of
2 (Pribyl, 2010) to convert from SOM to soil carbon, that is, SOM
is 50% car bon, and a liter ature value for s oil density of 144 0 kg/
m3 for sandy loam (Yu et al., 1993).Above-grounddr ybiomasswas
estimated for the meadow by counting the hay harvest in bales,
weighing a bale, calculating the proportion of water in a bale by oven
drying a sample and multiplying up. These values are not included in
the carbon sequestration figures as the pool is short lived; neverthe-
less, the productivity of the meadow is noted here.
2.3.3 | Relativereflectance
Albedoiscalculatedfromtheratioofreflected light to down-coming
light on a vertical surface. Here, we measure relative reflectance with-
out vertical surface images or down- coming light estimation, as a proxy
for albedo. Relative reflectance values were calculated for meadow and
lawn from images at three times of year: a March 2020 satellite image
retrieved from zoom.earth, an aerial view of the meadow in early flower
26 May 2020, and a phone camera photograph taken on the ground
(October20).AnalysiswasperformedinImageJ(Gilchrist,2011).
2.4 | Society
Asurvey was designed to assess respondents' opinions of the cul-
tural services provided by meadow and lawn, and respondents' pref-
erencesformeadowandlawn(AppendixS1). Ethics oversight for the
survey design and administration was provided by the Cambridge
Hub. The sur vey was administered once in 2021 with responses re-
cordedbetween6Febr uar yand26March.Atthist ime,themeadow
had had one flowering season and was in a winter dormant period.
Given the timing and method of recruitment respondents are likely
to have seen the meadow for themselves, although we did not insist
onthis.Atotalof278respondentswererecruitedviamailinglistsof
the University faculties, colleges, societies and University- affiliated
organisations. Respondents were informed of the purpose, methods
and end use of the research and gave their informed consent to their
data being collected and used for the purposes described in a privacy
notice.Anopt-outofhavinganswersquotedwasprovided.Norisks
to participants were identified and participant s were free to with-
drawatanytime.Asmallfinancialincentivewasof feredtorespond-
ents in th ef ormofa nA mazongiftvo ucherawar de dtot wor an domly
chosen respondents. Participants remained anonymous, unless they
opted into be ing contacte d for the randoml y selected rew ard. All
identifying information was deleted after disbursement of the re-
wards. Questions were always asked in the same order. The ques-
tion of preference for lawn, meadow or a mixture was repeated after
the provision of information on the benefits of lawns and meadows.
This information consisted of a written summary of the provisioning,
regulating, cultural and supporting ecosystem services derived from
wildflower meadows and lawns and was writ ten by the survey ad-
ministrator from published peer- reviewed literature. References to
the primary sources were provided to participants. Responses were
analysed using Wilcoxon signed- rank tests and chi- squared tests of
association. Open responses were analysed by identifying and ex-
ploring common themes qualitatively.
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3 | RESULTS
3.1 | Biodiversity
3.1.1 | Speciesrichness,abundanceandbiomass
Plant species richness was 3.6 times higher in the meadow compared
with its lawn control in 2021 (means 15.8 vs. 4.4 species; t = 8.497,
df = 4.773, p < 0.001) (Table 1). The combined spider and bug species
richness recorded during sweep netting was 3.7 times higher in the
meadow than the lawn (2021, average 6.6 vs. 1.8 species t = −4.382,
df = 5.039,p = 0.007),while the number of spiderandbugindividuals
caught in pitfall traps was 3.8 times higher in the meadow than the
lawn (2021, 7.6 vs. 2 individuals, respectively; t = 3.572, df = 6.23,
p = 0.011)( Table 1). At the top of the f ood chain, in sectivoro us bats
were recorded 3.1 times more often over the meadow than over the
lawn (100 vs. 32 tim es, respectively; pair ed t = 5.249,df = 21,p < 0.001),
and on average more bat species were recorded on the meadow each
night (4.31) compared with the lawn (3.45) (paired t = 2.425, df = 21,
p = 0.024)(Table 1). Total invertebrate biomass was 25 times higher in
the meadow than in th e lawn (2021, t = −6.7757,df = 4.2247,p = 0.002).
The mean b ody length of law n arthropod s pecies was 4.79 mm, c f.
mean body length of 8.75 mm for meadow arthropods (t = −3.78,
df = 41.5,p = 0.0005).Below-ground,therewasnodifferenceinnema-
tode species richness nor abundance in the meadow cf. lawn in 2021,
nor in the meadow in 2021 compared with its lawn baseline in 2019.
Meadow biomass was estimated from bale production; 142 bales came
off the meadow in 2020 versus 322 bales in 2021, giving an estimated
productivityof2.12and5.07 Mg /ha/year,respectively.
During BACI hy pothesis te sting (Table 2), a significant interac-
tion term was found for plant species richness, with the meadow
plant richness significantly higher than its original baseline and the
lawn control in all years (Figure 2; Table 2; Table S1 for contrasts). For
bats, abundance increased over the growing season more sharply
over the meadow than over the lawn; by October the meadow had
been mown, and there was no dif ference in bat activity at this time
point. For nematodes, abundance was significantly increased in
2020 in both the meadow and lawn relative to both 2019 and 2021
and was significantly higher in the meadow than the lawn in 2020
only. Thus, the greater abundance in 2020 was not sustained into
2021, and being an intermediate year, it does not feature in the
(non- significant) comparisons tested in Table 1. Nematode richness
was significantly higher in 2021 for both meadow and lawn com-
pared with 2020 and 2019, that is, although there were more nem-
atode genera in total in 2021, there was no difference between our
meadow and lawn treatment of interest (as is tested in Table 1).
Atotalof84plantspecies,16bugandspiderspecies,149nema-
tode genera and 8 bat species were recorded during sampling. Of the
84 plant species recorded in 2021, only 33 were sown species (24
from the perennial meadow species seed mix, 6 persisting from the
cornfield annual mix and 4 of the supplementar y mix: Odontites ver-
nus, Onobrychis viciifolia, Ononis spinosa and Iberis amara, a nationally
sc ar cesp ecies) .A dd ition al ly, ni ne sp eciesofpara si ticmicrofun gi(suc h
as powdery mildews and rusts) were recorded by C. Preston on three
visits in June, July andAugust 2022, including a new county record
of the fungal plant pathogen Cercospora zebrina on Medicago arabica.
3.1.2 | Speciesofconservationpriority
The meadow suppor ts 14 species with conservation designations,
compared with 6 species with conservation designations in the
lawn.Allthedesignatedspeciesrecordedin the lawn werealsore-
corded from the meadow (Lygus pratensis, Barbastellus barbastellus,
FIGURE 1 King'sBackLawnand
meadow (photo 10 June 2020, ©Geoff
Robinson/BAVMedia).The10sampling
locations for plants and invertebrates,
and two sites of static bat detectors, are
marked. The land to the west of the river
is Scholars' Piece. The white building
facing the lawn is the Gibbs building. The
Chapel stands to the north of the Gibbs
building.
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Eptysicus serotinus, Nyctalus leisleri, Pipistrellus pipstrellus), apart from
Pipistrellus nathusii, which was not recorded over the meadow. The
full list of meadow species with conser vation designations has nine
plants ( Agrostemma githago, Glebionis segetum, Bupleurum rotundifo-
lium, Euphrasia confusa, Iberis amara, Knautia arvensis, Ononis spinosa,
Briza media, Onobrychis viciifolia), four bats (Barbastellus barbastellus,
Eptysicus serotinus, Nyctalus leisleri, Pipistrellus pipstrellus) and one
bug (Lygus pratensis). Euphrasia confusa (det. J. Shanklin 2020) had
not previously been recorded in Cambridgeshire.
In 2021, the meadow supported approximately four times as
many declining plant species (those with negative change indices)
compared with the remaining lawn (44 cf. 11) and the original lawn
baseline (44 cf. 10). Meadow plant species were on average rarer
than lawn species, with meadow species found in 1982 hectads
(10 km × 10 km) across GB, c f.246 3 hectads for t he lawn species.
Meadow spiders had a non- significantly slightly wider distribution
than lawn species (means 1255, 1219, n = 17,14), and have shown
a greater increase in records post- 1992 (+99% cf +73%). Rarer bugs
included Dufouriellus ater (28 hectads, meadow) and Lygus pratensis
(92 hectads, meadow and lawn). No data on nematode species' con-
servation status or range have been published to our knowledge.
3.1.3 | Speciescompositionandhabitatassociations
Samples are ordinated in Figure 3. For plants (Figure 3a), axis 1 dis-
tinguishes clearly the two meadow habitat s (meadow 2020, meadow
2021) from the lawn habitats in all years, indicating different species
composition between the habitats. The three lawn years and the
pre- sowing meadow baseline 2019 (=lawn habitat) are not distin-
guishablefromeachother.Axis2distinguishesthemeadowin2020
from the meadow in 2021, as floral composition shifts year on year.
For spiders and bugs from the sweep samples (Figure 3b), axis 1 also
separates meadow habitat from lawn habitat ver y clearly, while axis
2 separates both habitats by year, showing that species composition
is influenced by both habitat type and annual variation. For nema-
todes (Figure 3c), samples are grouped by year, rather than habitat
type, suggesting that interannual variability in samples is the most
significantinfluenceonspeciescomposition.Axis1separates2021,
while axis 2 separates 2019 from 2020. Habitat types are barely
distinguishable within years. For bats (Figure 3d), axis 2 separates
samples clearly by habitat, indicating different species composition
betweenthehabitats.Axis1separatesthreesamplesfromtherest,
nights on which a lot of bat activity was recorded.
TAB LE 1 Meanrichnessandabundanceofplants,spidersandbugs(pit falls,sweeps),nematodesandbatsrecordedinthemeadowin
2021, compared with the presowing meadow baseline (2019) and lawn control (2021). Significance tests are Wilcoxon signed- rank tests,
paired for bats only. The change in each response variable is reported first. Significant changes are shown in bold, ***p < 0.001,**p < 0.01,
p < 0.05*.Checklisttotalsareincludedattheendofthetable.
Response variable Meadow 2021
Presowing baseline
(meadow 2019)
Change ratio cf
baseline Lawn 2021
Change ratio cf
control
Plant mean richness 15.8 4.2 3 .76* *
W = 25,p = 0.0097
4.4 3.59*
W = 25,p = 0.011
Plant mean abundance 91 77. 8 1.17
W = 18,p = 0.31
62.6 1.45
W = 21,p = 0.095
Pitfall mean richness 33.2 0.94
W = 14.5,p = 0.32
1.75 1.71
W = 11.5,p = 0.91
Pitfall mean abundance 7.6 51 .52
W = 18,p = 0.29
23.8*
W = 19,p = 0.032
Sweep mean richness 6.6 N/A N/A 1.8 3.67*
W = 25,p = 0.011
Mean inver tebrate biomass
(pitfalls) (g)
5.60 N/A N/A 0.225 24.89*
W = 20,p = 0.016
Nematode generic mean
richness
48.2 31.4 1.54
W = 13.5,p = 0.92
49. 2 0.98
W = 20,p = 0.15
Nematode mean abundance 116 9.4 2805.8 0.42
W = 4,p = 0.095
970.4 1.21
W = 15,p = 0.69
Bat mean richness 4.31 N/A N/A 3.45 1.25*
V = 134.5,p = 0.032
Bat mean abundance 100.32 N/A N/A 32.36 3.10***
V = 224,p = 0.00017
Tot a ls : plan t s 84 22 3.82 28 3.00
Totals: pitfalls 710 0.70 71.00
Tot a ls : swe eps 16 N/A N/A 91.78
Totals: nematode genera 90 54 1.66 89 1.01
Tot a ls : bat s 8N/A N/A 71.14
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TAB LE 2 Before-after-control-impact(BACI)hypothesistestingwithANOVA.Contrastsarecalculatedpost-hocusingTukeyHSD
(Table S1).
df F p Direction of significant contrasts
Plant richness, ln transformed
Location (lawn, meadow) 1,24 1 28.13 <0.0001 Meadow > lawn
Year (2019, 2020, 2021) 2,24 39.92 <0.0001 2021 = 2020 > 2019
Location × year 2,24 19. 24 <0.0001 Meadow:2020 > lawn:2019
Meadow:2021 > lawn:2019
Meadow:2020 > meadow:2019
Meadow:2021 > meadow:2019
Meadow:2020 > lawn:2020
Meadow:2021 > lawn:2020
Lawn:2021 < meadow:2020
Meadow:2021 > lawn:2021
Plant abundance, untransformed
Location (lawn, meadow) 1,24 8 .705 0.00663 Meadow > lawn
Year (2019, 2020, 2021) 2,24 1.724 0.20066 N /A
Location × year 2,24 0.049 0.8270 0 N/A
Pitfalls richness, untransformed
Location (lawn, meadow) 1,22 2.357 0 .139 N/A
Year (2019, 2020, 2021) 2,22 0.645 0.535 N/A
Location × year 2,22 0.226 0.800 N /A
Pitfalls abundance, untransformed
Location (lawn, meadow) 1,22 10. 5 41 0.0037 Meadow > lawn
Year (2019, 2020, 2021) 2,22 1.054 0.3654 N/A
Location × year 2,22 1.818 0 .1859 N/A
Sweep richness, ln transformed
Location (lawn, meadow) 1,16 54 .418 <0.0001 Meadow > lawn
Year (2020, 2021) 1,16 5.19 9 0.35800
Nematode richness, ln transformed
Location (lawn, meadow) 1,24 0 .270 0. 60791 N /A
Year (2019, 2020, 2021) 2,24 8 .922 0.0 0127 2021 > 2020 = 2019
Location × year 2,24 0.111 0. 89498 N/A
Nematode abun, sqrt transformed
Location (lawn, meadow) 1,24 5 .172 0.03219 Meadow > lawn
Year (2019, 2020, 2021) 2,24 7 7. 3 8 6 <0.0 005 2020 > 2019 > 2021
Location × year 2,24 8.250 0.00188 Meadow:2020 > lawn:2019
Meadow:2020 > meadow:2019
Lawn:2021 < meadow:2020
meadow:2021 < meadow:2020
Lawn:2021 < lawn:2020
Meadow:2021 < lawn:2020
Meadow:2020 > lawn:2020
Lawn:2020 > meadow:2019
Lawn:2020 > lawn:2019
Bat richness, untransformed
Location (lawn, meadow) 1,39 3.804 0.05834. Meadow > lawn
Month 3,39 6.509 0.00112 May = June < July < October
Bat abundance, sqrt transformed
Location (lawn, meadow) 1,36 24.534 <0.0001 Meadow > lawn
Month 3,39 8.422 0.000227 May = June < July < Oc tober
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df F p Direction of significant contrasts
Location × month 3,36 2.929 0.046653 Meadow:June > lawn:May
Meadow:July > lawn:May
Lawn:October < meadow:May
Meadow:June > lawn:June
Meadow:July > lawn:June
Lawn:October < meadow:June
Meadow:October < meadow:June
Meadow:July > lawn:July
Lawn:October < meadow:July
Meadow:October < meadow:July
TAB LE 2 (Continued)
TAB LE 3 Emissionsandcost sassociatedwiththemowingandfertilisingregimesofmeadowandlawnplantings,thet womostcarbon
intensive management activities, for King's Back Lawn, and Cambridge College green space in total. See Table S2 for constituent values and
references.
Lawn Meadow
GHG emissions (Mg CO2- e/
ha/year)
Co st (GB P/ha/
year)
GHG emissions (Mg CO2- e/
ha/year)
Co st (GB P/
ha/year)
Mowing 0.891 659. 62 0.0122 9. 0 4
Fertiliser 0.484 532.80 0 0
Mowing + Fertilising 1.375 119 2. 42 0.0122 9.0 4
TotalforKing'sBackLawn(0.99 ha) 1 .3 61 1180.49 0.0120 8.95
Total lawn across all Cambridge
colleges(4 3.7 ha)
60.08 52108 .6 4 0.53 394. 87
FIGURE 2 Speciesrichnessandabundancefora = plant s,b = sweeps(spidersandbugs),c = pitfalls(spidersandbugs),d = nematodes,
e = bats.Confidenceintervalsaremean ± (1.96 × SE).Abundancedatawerenotcollectedduringsweepnetting(panelb).Abundancevalues
are × 1000yaxisvaluesinpanel(d).Verticalbluelinebetween2019and2020denotesmeadowestablishment.
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For plants in 2021, no association between broad habitat
preferences and location was found overall (X2 = 12.706, df = 12,
p = 0.3908). Most strongly overrepresented are improved grassland
speciesinthelawn(broadhabitat5,standardisedresidual = 1.74)and
calcareous grassland species in the meadow (broad habitat 7, stan-
dardisedresidual = 2.54).Thespidersfoundinoursurveyarehabitat
generalists, especially of grasslands and gardens, with a few of the
meadow species characteristic of warm, sunny & exposed situations
(Enoplognatha latimana, Xysticus kochi, Ozyptila sanctuaria), which are
also the scarcer of the species being generally restricted to the south.
Allsevenbugspeciesofthelawnwerealsofoundinthemeadowex-
cept Scolopostethus thomsoni, a generalist species feeding on nettles.
Bug species of the meadow are grassland species or generalists, with
most notable species including Lygus pratensis, a meadow specialist
mirid bug with a southern but expanding distribution and Orthops kal-
mii and O. campestris, both umbel- feeding mirid bugs abundant in July,
when the carrots Daucus carota were the dominant meadow plant.
Nematode functional guilds are defined as the combination of
nematode feeding habit and coloniser- persister (cp) classification.
The Enrichment Index and the Structure Index, both derived from
the weighted relative numbers of each of the functional guilds in
a sample, are descriptors of food web condition reflecting nutri-
ent status and maturity of habitat, respectively (Ferris et al., 2001).
Most samples fall within quadrant B, typical of managed grassland
and agricultural systems (Berkelmans et al., 2003) (Figure 4). These
have a high structure index value indicative of low disturbance or
undisturbed soils and a high enrichment index value symptomatic of
N- enrichment. The single exception is the meadow habitat in 2020:
FIGURE 3 NMDSordinationsofBray–Curtisdissimilarit ymatricesforsamples,t woaxeswerespecifiedwitheachordination(threeaxes
for bats) and all ordinations converged with stress <0. 2. a: plants, b: sweeps (spiders and bugs), c: nematodes, d: bats. Pit fall data were too
few for valid ordination.
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with a lower enrichment index (lower fertility), it just falls within
quadrant C, typical of natural grassland systems. However, this re-
sult is not recovered in 2021 (with the meadow back to quadrant B),
so this is not likely to be significant. Nematode func tional composi-
tion is similar across lawn and meadow.
For bats, there is a ver y significant association between spe-
cies and location overall (X2 = 128.21, df = 8 , p < 0.001; Figure 5).
Most species were associated strongly with the meadow (serotine
Eptesicus serotinus, common pipistrelle Pipistrellus pipistrellus, brown
long- eared Plecotus auritus) or lawn (leisler's Nyctalus leisleri, noct-
ule Nyctalus noctula, nathusius' pipistrelle Pipistrellus nathusii), while
barbastelle Barbastella barbastellus, daubenton's Myotis daubentonii
and soprano pipistrelle Pipistrellus pygmaeus were associated with
each habitat equally.
3.2 | Climate change
3.2.1 | Emissionsreductionsandcostsavings
Using literature values and the maintenance regime at King's, we
estimate that greenhouse gas emissions (CO2- e/ha/year) are 112
times higher from lawn than meadow, while the maintenance re-
gime is 132 times costlier for lawn than meadow ( Table 3).Across
all Cambridge colleges, converting all the current lawn area of
43.7 hato meadowplantingwouldreduceannualgreenhousegas
emissions by 59.55 Mg CO2- e each year, and cost £51,713 less
eachyeartomaint ain.Overall,weestimateemissionsof1.375 Mg
CO2- e/ha/year from lawn (of which 65% came from mowing and
35% from fer tilising) ver sus 0.0122 Mg CO2- e/ha/year from the
meadow forasingle mow,giving a saving of 1.36 Mg CO2- e/ha/
year. King's has made available to other colleges and the Cit y
Council hay bales for green haying, eliminating the cost of seed,
although there would still be site preparation costs like scarif ying
to consider. The area of greenspace managed by each college is
estimated in Figure S3.
3.2.2 | Carbonsequestration
In 2020, with 10 samples (mean of three nested pseudoreplicates),
we found no statistically signific ant difference in the mean SOM
(SOM%) of the formal lawn (16%) and wildflower meadow at King's
(17.3%) (t = −1.44, df = 5.69, p = 0.2). Soil carbon was estimated at
124.56 Mg/h a in the top 0–10 cm. T he carbon po ol stored by the
plant below- ground, for example, in root s was not measured.
FIGURE 4 Functionalecologyofthenematodecommunities
derived from the weighted relative numbers of each functional
guild in the samples. Enrichment index is a descriptor of food web
condition reflecting nutrient status with higher scores having
higher nitrogen availability. Structure index describes maturity of
thehabitat,withhigherscoresbeinglessdisturbed.Orange = lawn,
blue = meadow;squares = 2019,circles = 2020,triangles2021.
FIGURE 5 Batspecieshabitatassociations.Coloursandindexrefertothestandardisedresidualsofachi-squaredtest,X2 = 128.21,df = 8,
p < 0.001.BARBAR = barbastelleBarbastella barbastellus,EPTSER = serotineEptesicus serotinus,MYODAU = daubenton'sMyotis daubentonii,
NYCLEI = leisler'sNyctalus leisleri,NYCNOC = noctuleNyctalus noctula,PIPNAT = nathusius'pipistrellePipistrellus nathusii,PIPPIP = common
pipistrelle Pipistrellus pipistrellus,PIPPYG = sopranopipistrellePipistrellus pygmaeus,PLEAUR = brownlongearedPlecotus auritus.
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3.2.3 | Relativereflectance
Analysi s of a series of photo graphs incl uding both the m eadow and
lawn, ta ken throughout th e year, showed the meadow re flectance to b e
25%– 34% higher than that of the adjacent lawn (Figure S2; Table S4).
3.3 | Society
Our attitudes survey attracted 278 respondents, a mixture of un-
dergraduate students (50%), postgraduate students (23%), non-
academic staf f (13%), academic staf f (10%), alumni (2%) and those
with no affiliation (2%). 83% of respondents had a college affilia-
tion, with 31 colleges represented. Sur vey questions can be found
inAppendixS1.
The response to meadows was overwhelmingly positive.
Respondents considered meadow more aesthetically pleasing
than lawn (mean score 9.22 cf. 6.53, Wilcoxon W = 10,661, p-
value < 0.0001)andmoreenvironmentallyfriendlythanlawn(mean
score 8.90 cf. 3.77, Wilcoxon W = 2068, p-value < 0.0 001). 68% of
respondents said they would prefer a mixture of lawn and meadow
on campus/in their area, 30% said they would prefer entirely
meadow to lawn, and just 1.4% (4 respondents) preferred entirely
lawn. Significantly more respondents (83.5%) reported that mead-
ows suppor ted their mental well- being (‘Meadows are great, they
heal my soul ’; ‘Meadows would dramatically improve my well- being’),
compared with lawns (45%) (X2 (275,278 ) = 89.6, p < 0. 001). 38.1%
of respondents thought meadows had educational value, compared
with just 4% for lawns. However, more respondents thought lawn
provided recreational services (76%) than thought meadow provided
recreational ser vices (56%). Af ter beingpresented with written in-
formation about the benefits of both lawn and meadow, 52% said
they preferred a mix ture of lawn and meadow, and 47% said they
preferred entirely meadow to lawn (and 1% still preferred lawn),
that is, respondents tended to move from a position of ‘mixture’ to
‘entirely meadow’. There was no interaction between affiliation and
habitat preference (X2 = 9.49,df = 10,p = 0.49),nor collegeand hab-
itat preference (X2 = 55.7,df = 62,p = 0.70).ThefourKing's respon-
dents preferred meadow (2) or a mixture of lawn and meadow (2).
In Cambridge, most colleges restrict access to their lawns to se-
nior members of the college only, that is, access is denied to student s
and visitors. This reduces the amenit y value of lawn in collegiate
Cambridge, while also increasing the perceived status of those
lawns. Respondents were then divided as to whether that makes
lawns a problematic emblem of classism and misdirected author-
ity, or a heritage aesthetic that ought to be retained. For example,
‘Lawns are a symbol of elitism and exclusion at Cambridge’. ‘I find the
overly tended lawns of the Cambridge colleges sterile and uninviting
(not helped by the fact that one is often told not to walk on them)’.
‘I find the lawns at Cambridge to be, in general, quite stuffy, clas-
sist, off- putting and generally self- defeating. (What is the point of
grass you can't walk or sit on?)’. ‘Lawns just seem really pretentious
to me’. ‘Particularly in Cambridge, lawns are forbidden territory’. ‘I
really like meadows and I understand that they are better environ-
mentally (diversity of species etc), but I also really like to be able to
sit on grass, picnic with friends, kick a football with children (obvi-
ously I'm not talking about the posh Cambridge lawns that you're
not allowed to walk on!)’. ‘I think that some of the grass courts in
Cambridge are sor t of iconic, so it'd be a shame to lose them all.
That said, I'd definitely prefer the majority of green space in my col-
lege to be meadow’. ‘I would only want a lawn over a meadow if I
was allowed to use the lawn for recreational purposes. This is not al-
lowed in most Cambridge colleges so I see no benefit to a lawn over
a meadow’. ‘I don't think the characteristic square/rectangular lawns
of Cambridge colleges would look good as meadows’.
When respondent s were asked if they had any objections to
the conversion of lawn to meadow, 66% of 175 respondents had
none at all. Recorded objections were that lawn area for recreation
should be maintained e.g. for sports and sitting (22 respondents),
aesthetic concerns (12 respondents), concern over increased
hayfever, insect stings and ticks (8 respondents), that accessibil-
ity needed to be maintained (5 respondents), that species choice
should be sensitive, i.e. native, low water demands, local prove-
nance, good for pollinators (4 respondents), that it could cause
controversy (2 respondents), that meadows look messy in winter
(2 respondents) and that disruption during the conversion should
be minimised (1 respondent). We asked respondents to suggest
alternatives to replacing lawn with wildflower meadow with the
following response rate: herb lawn, moss lawn or living lawn (8),
woodland (8), trees (8), no maintenance of current lawns (7), allot-
ments (6), ponds (5), formal garden (3), flower border (2), scrub (2)
and corn maze (1). Incidentally, Trinity College had a maze on it s
grounds in 1592 (Willis & Clark, 1886).
When asked if meadow planting could contribute to the
University's sustainability goals, 164 of 170 respondents thought
yes (96%). Of the six respondents who said no, two said meadow
planting was tokenism only, and four did not elaborate. One said ‘re-
placing a tiny area of ornamental lawn in Cambridge will not affect
UK biodiversity in a serious way— there are better hills to die on’.
Afew who thoughtthatmeadow couldcontribute tosustainability
goals also worried about tokenism, ‘taking par t in such a visible move
towards sustainability will let colleges off the hook for more diffi-
cult and meaningful change and practices’. Thirteen respondents
made comments around the positive impact being modest only (e.g.
‘maybe on a local level’, ‘not as much as other interventions’, ‘bigger
fish to fry’). On the other hand, ‘huge impact’ ‘big impact’ ‘Yes!’ ‘defi-
nitely’ ‘much more’ ‘absolutely’ ‘of course’ ‘obviously’ and ‘I'm sure’
appeared in 18 responses.
Twenty respondents highlighted meadow planting as an im-
portant sustainability leadership action that the University should
take. ‘The King's meadow is a ver y visible sign of positive change’.
Respondents commented that planting meadows ‘could change
mindset about sad lawn culture’ and would be ‘a strong message
that the universit y realises the importance of biodiversit y in cre-
ating resilient ecosystems and preventing extinction’. One com-
mented, ‘The lawns are symbolic in many ways of Cambridge's
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link to tradition, and if this small element of tradition changes I am
hopeful that traditionwon'tbesucha barrierelsewhere’.Another
thoughtthat‘AsaleadingUniversity,Cambridgehasaresponsibil-
ity, not only to the environment but also as an example to other uni-
versities to make biodiversity an important issue’. One respondent
summarised, ‘We need to abandon pointless traditions like lawns”,
while another had a simple message for us: ‘Meadow good lawn bad
save the planet’.
4 | DISCUSSION
4.1 | Value to wildlife
In spite of its diminutive dimensions and recent establishment,
the meadow is of considerable and demonstrable value to wild-
life. Compared with the remaining lawn, the meadow suppor ts ap-
proximately three times more plant species and a much taller sward,
which in turn supports three times more spider and bug species and
individuals, while terrestrial invertebrate biomass is a massive 25
times higher in the meadow compared with the lawn. Bats were re-
corded three times more often over the meadow than the lawn. The
meadow is at tracting s pecialist spe cies, like Lygus pratensis, a meadow
specialist mirid bug with a southern but expanding distribution, and
Orthops kalmii and O. campestris, both umbel- feeding mirid bugs
abundant in July when the wild c arrots were the dominant meadow
plant. The weekly moth trap we run at the King's meadow also at-
tract s a disproportionate number of grassland- specialist moth spe-
cies, including rarities like Noctua orbona (Lunar Yellow Underwing),
and meadow specialists like Bucculatrix nigricomella (Daisy Bent-
wing) (Marshall et al., 2022). Fourteen species with conservation
designations were recorded in the meadow, more than twice that
of the lawn. There are four times more declining plant species in the
meadow than the lawn and two new records for C ambridgeshire: the
eyebright species Euphrasia confusa, and the microfungus Cercospora
zebrina on Medicago arabica. The decline in once- common arable
weeds across the UK has been accompanied by a concomitant de-
cline in their obligate fungal parasites (Preston, 2022). Of note at
the King's meadow is Puccinia cyani, a rust species of cornflower
Centaurea cyanus once considered ‘extinct in the UK along with its
host’ ( Termorshuizen & Swertz, 2011). This species, along with the
powdery mildew Peronospora agrostemmatis (host plant corncockle
Agrostemma githago), now seems to be enjoying a resurgence in wild-
flower mixes in Cambridgeshire and beyond (Preston, 2022).
Plant species composition and sward height were both found to
be impor tant drivers of invertebrate abundance in grassland sys-
tems, with most inver tebrate orders found to be more abundant
where vegetation height was longer than mown grassland (Norton
et al., 2019).AtKing's,invertebratebodylengthwasonaveragelon-
ger in the meadow than the lawn, and we suggest that the increased
sward height of the meadow allows larger- bodied taxa to avoid bird
predation, increasing all of species richness, abundance and biomass
(Figure S1).
Previous studies have shown that plant diversity has a strong
bottom- up effect on multitrophic ecosystems connected in food
webs, with particularly strong effects on lower trophic levels like
herbivores (Scherber et al., 2010).At King's, insectivorousbatsat
the top of the food chain were recorded three times more often
over the meadow and the lawn, which we attribute to increased
foraging activity associated with higher invertebrate biomass. The
proximity of the river corridor probably increases the conspicuous-
ness and at trac tiveness of the meadow to bats in compensation
for its small size. The bottom- up control of restored species- rich
meadows apparent at King's is encouraging from an urban resto-
ration perspec tive, as it is relatively straightforward to manipulate
plant species richness and sward height to give positive effects for
other taxa.
The size of habitat patches and their connec tivit y have been
identified as key to maintaining high levels of urban biodiversit y
(Beninde et al., 2015). In that meta- study, the smallest area consid-
ered necessary to sustain species numbers before they decreased
exponentially was 1 ha.Asmall area (1.7 ha) ofwildflowermeadow
had significantly higher insect abundance and species richness than
a comparable amenity grassland (Hutchinson et al., 2020). At just
0.36 ha, thecurrentmeadow extent at King's is much smaller than
is typical for previously studied grassland systems, although even
‘mini-meadows’ ofjust 2 × 2 m have been shown to have a positive
impact on wildlife (Griffiths- Lee et al., 2022). To have recorded such
dramatic and positive changes for biodiversity, across trophic levels,
from so sma ll an urban area is hi ghly encouragi ng for future urb an res-
torationprojects.Connectivitydependsonthetaxonunderstudy.A
single night of small mammal trapping at the meadow in 2021 caught
no animals, while one wood mouse was caught at Scholars' Piece.
For small mammals, the meadow is probably a relatively isolated and
inaccessible island of habitat, bounded by Clare College, the Gibbs
building, the river and other college buildings. Bats and aerial insects
on the other hand, including grassland specialist species, seem to
have dispersed to make use of the area without problems.
Four of every five multicellular animals on the planet are nema-
todes, and their sensitivity to pollutants and environmental distur-
bance makes them an excellent indicator taxon for studying changes
to ecosystems below- ground (Bongers & Ferris, 1999). At King's,
there is no difference in nematode species richness, abundance nor
functional guild composition in the meadow cf. lawn. The functional
guild composition has values t ypical of managed grassland and agri-
cultural systems (Berkelmans et al., 20 03). Changes to below- ground
fauna are expected to be slower than for above- ground fauna: dif-
ferences in nematode communities along plant species diversity
gradie nts have been r eported a fter 15 year s (Dietrich e t al., 2021)
and after12 years (Viketoftetal., 2009). Over a seven- year study,
nematode diversit y was little influenced by plant diversity, and
the faunal composition did not stabilise but changed continuously
(Sohlenius et al., 2011). Our nematode datasets show similarly high
heterogeneity in richness, abundance and to a lesser extent compo-
sition year- to- year and sample- to- sample. While sequencing costs
prohibit denser sampling, inherent soil heterogeneity seems to have
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MARSHALL et al.
precluded the detection of any compositional changes in the early
stages of meadow establishment, if they were there.
4.2 | Climate change mitigation
Reduced maintenance and fertilising associated with the meadow
reduced e missions by 1. 36 Mg CO2- e per hectare per year com-
pared with lawn. These estimates are similar to previous estimates
of1.0 and 1.6 Mg CO2e/ha/year for turf on Swedish golf courses
(Tidåker et al., 2017).Althoughtheabsolute CO2- e emissions re-
duction is small relative to the College's total emissions of around
1600 Mg CO2- e/year, it has been achieved with a cost saving of
approximately £650/ha/year, and increased value to both people
and wildlife.
Our small sample size did not reveal a significant difference in
soilorganiccarbonbetweenthelawnandmeadowafter2 years,at
least not in thetop10 cm. Deeperrooted species in the meadow
compared with lawn may increase soil carbon <10 cm. Dete cting
changes in soil carbon stocks even over a 5- year period is ver y
difficult, unless the carbon input change to the soil is very great
(>20%) or the sampling density is ver y high (Smith, 2004). Our es-
timateoftotalmeadowsoilcarbonstockof123.55 MgC/hainthe
top0–10 cmishighcomparedwithliteraturevaluesforcalcareous
grasslands and even higher for horticultural soil. Typical soil car-
bonstocksareapproximately69 MgC/hatoa15 cmdepth(C arey
et al., 2008)or approximately51 MgC/hatoadepthof10 cm for
upland calcareous grazing pasture (Niklaus et al., 2001). While we
have no reason to doubt the reliability of our relative estimate of
SOM concentration between meadow and lawn samples, our total
estimate of soil carbon stock is likely prone to substantial measure-
ment error associated with assumptions made for soil bulk density,
organic matter- carbon conversion factor and stone content (Gregg
et al., 2021). Under particular conditions, lawn may act as a net
sinkofatmosphericcarbon( Velasco etal., 2016). Over the whole
season, biogenic CO2 fluxes from soil respiration far exceeded an-
thropogenic fluxes associated with mowing and other management
(Lerman & Contosta, 2019). Soil CO2 flux is, thus, an important part
of the overall carbon budget of grassland and one that we have not
studied at King's.
Reflectance increased by 25%– 34% for meadow relative to
lawn. Unmown lawn also had a higher reflectance (+25%) t han
formal lawn (data not presented), suggesting maintenance regime
rather than species composition may be the key factor controlling
reflectance. The meadow is dominated by white- flowered species
throughout the flowering period (Austrian chamomile Anthemis
austriaca, followed by ox- eye daisy Leucanthemum vulgare and
carrot Daucus carota), although higher meadow reflect ance was
alsoobservedduringnon-floweringperiods.Alpinemeadowplots
with flowers removed manually had lower albedo and resultantly
warmer soil temperatures than those with typical floral density
(Iler et al., 2021).AstheCambridgeColleges'lawnareais47.3 ha,
or 1.3% of the city of Cambridge, relaxing the maintenance regime
of formal lawns or replacing some of the plantings with meadow
could increase reflectance and thus albedo, helping to maintain
a cooler urban microclimate under future global warming (Yan
et al., 2019).
4.3 | Society
Our survey respondents thought meadows provided greater aes-
thetic, educational and mental well- being services than lawns, as
well as providing a stronger cultural connection, having higher
religious/spiritual value, and being more inspirational than lawns.
Meadows scored lower than lawns for recreational and practi-
cal services, emphasising the need for heterogeneity of plant-
ing within managed urban ecosystems. Access to green space
has value for our health, mental (Fuller et al., 20 07) and physi-
cal (Mitchell & Popham, 2008). The latter study demonstrated
how income- related inequality in health was less pronounced
where there was greater accessibility to green space, pertinent
considering Cambridge is the most unequal cit y in the United
Kingdom (CfC, 2018). Our respondents, who were overwhelm-
ingly members of the collegiate university and mainly students,
were acutely attuned to signals of land ownership and stew-
ardship and the issue of unequal access to greenspace, themes
that occurred frequently in open answers without prompting. It
is relatively uncommon to accommodate hundreds of people in
communal facilities where access to greenspace is so explicitly
socially hierarchical and exclusionary. The strong support for
meadow amongst our surveyed population may well be attribut-
able in par t to dissatisfaction with the current access to greens-
pace arrangements amongst the surveyed community and would
not necessarily be expected amongst the city population overall,
orindifferentcities.Additionally,theimmensepopularityofthe
meadow planting apparent in this cohor t may not persist over
time, if residents come to see this habitat as typical and, thus, less
associated with novelty and action for the environment (Bullock
et al., 2021). Where lawns were once used to demonstrate how
much land one could keep out of productivity, it seems that they
still have a symbolic significance around stewardship today, with
respondents agreeing that meadow planting can be used to signal
sustainability leadership.
Restoration and conservation efforts, especially within urban
areas, should be a give- and- take between what is best for wildlife
and what is best for people. Our survey showed that the local com-
munity would prefer a mixture of lawn and meadow, compared with
either lawn or meadow monoculture, with only 1.4% preferring en-
tirely lawn. More respondents found meadow or meadow- mix plant-
ing favourable when the option of paths and seated areas was added.
Similar responses were recorded in Sweden, where residents valued
lawn for playing, resting, picnicking, walking and socialising but overall
preferred a variet y of planted spaces that provided good conditions
for different senses (sound, smell, touch and sight) and a range of ac-
tivities (Ignatieva et al., 2017). In China, planted wildflower meadows
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MARSHALL et al.
have become a socially accepted landscape type, with respondents
preferring meadows with a colourful and natural appearance (Jiang
& Yuan, 2017 ).
5 | CONCLUSIONS
Establishing meadow in place of lawn at King's has improved wild-
life value and reduced greenhouse gas emissions, while providing
greater benefits to the college and wider Cambridge community at a
lower cost. Our biodiversit y study revealed that increased meadow
planting in place of lawn would increase the value of the colleges
and university estates to wildlife by about threefold. Wider meadow
planting would have a positive impact on climate change mitiga-
tion eff orts, savi ng around 1.36 Mg CO2- e per converted hectare
per year. Meadow planting would have additional climate change
adaptation benefits including increased albedo via higher reflec-
tances and a reduced urban heat island effect. The number of cut s
required per year to maintain a lawn is expected to rise as growing
seasons become longer under climate change (Sparks et al., 2007),
while deeper- rooted perennial forb species are likely to better toler-
ate an intensified drought regime in Cambridge. These benefits have
been achieved with a cost saving of approximately £650/ha/year.
The Cambridge community proved overwhelmingly in favour of in-
creased meadow planting on the college and University estates, with
respondents happiest with a proposed mix ture of hard- wearing lawn
turf alongside more ecologically valuable and beautiful meadow
planting. Respondents were clear that meadow planting should be
in conjunction with maintaining or increasing access to greenspace
for recreation, which is particularly salient for collegiate Cambridge
where access to lawn has traditionally been restricted to senior
membersoftheCollegesonly.Alongsideretaining lawn for practi-
cal recreation purposes, there was some support for maintaining the
heritage aesthetic of Cambridge quads and lawns in some iconic set-
tings, although no respondents suggested that King's should revert
its Back Lawn meadow planting to lawn.
TheUniversitylauncheditsBiodiversityActionPlaninNovember
2020 (Cambridge Green Challenge, 2020) and 2022, a Colleges
BiodiversityAudithasbeeninpreparation.Thesedocumentsdefine
a 10- year vision for biodiversity on the collegiate Cambridge es-
tates, with an implementation plan and data- driven targets, effor ts
to which the positive results reported here can lend strong support.
Cambr idge respondent s thought it was important that the Un iversity
and Colleges should take a leadership role in the stewarding of
their estates for nature and wildlife, supporting effort s by the City
Council. This study takes an interdisciplinar y approach, highlighting
the value of small projects to local biodiversity, cultural services and
carbon sequestration. Beyond C ambridge, an estimated 25% of the
UK's urban area is grassland, and 84% of people live in urban areas.
We suggest that meadow establishment in place of some lawn area
is a small but worthwhile, and easily scalable, contribution to CO2
emissions reductions, which brings substantial additional benefits
for both people and wildlife.
AUTHOR CONTRIBUTIONS
Cicely A. M. Marshall: conceptualisation, methodology, software,
formal analysis, investigation, resources, data curation, writing—
original draft, writing— review and editing, visualisation, supervision,
project administration, funding acquisition. Matthew T. Wilkinson:
conceptualisation, methodology, investigation, data curation, re-
sources. Peter M. Hadfield: conceptualisation, methodology, inves-
tigation, resources. Steven M. Rogers: methodology, investigation,
resources, data curation, writing— review and editing. Jonathan
D. Shanklin: methodology, investigation, data curation. Brian C.
Eversham: conceptualisation, methodology, investigation, data cu-
ration. Roberta Healey: methodology, investigation, data curation.
Olaf P. Kranse: methodology, investigation, data curation. Chris
D. Preston: methodology, investigation, data curation. Steven J.
Coghill: conceptualisation, resources. Karris L. McGonigle: con-
ceptualisation, methodology, investigation, data curation. Geoffrey
D. Moggridge: conceptualisation, resources. Peter G. Pilbeam:
conceptualisation, methodology, investigation, data curation, re-
sources. Ana C. Marza: conceptualisation, methodology, software,
formal analysis, investigation, data curation. Darinka Szigecsan:
methodology, investigation, data curation. Jill Mitchell: methodol-
ogy, investigation, data curation. Marcus A. Hicks: methodology,
investigation, data curation. Sky M. Wallis: methodology, investi-
gation, data curation. Zhifan Xu: methodology, investigation, data
curation. Francesca Toccaceli: investigation, data curation. Calum
M. McLennan: investigation, data curation. Sebastian Eves- van den
Akker: conceptualisation, methodology, software, formal analysis,
investigation, resources, data curation, writing— review and editing,
project administration, funding acquisition.
ACKNO WLE DGE MENTS
The authors acknowledge support from the Gatsby Charitable
Foundation and King's College for two studentships. Work at the
Crop Science Centre at the University of C ambridge is supported by
BBSRC grants BB/R011311/1, BB/N021908/1 and BB/S006397/1.
Cambridge Hub supported the inter view questionnaire design via
the Engage for Change programme. Cicely Marshall, Sebastian Eves-
van den Akker, Geoffrey Moggridge, Steven Coghill, Sk y Wallis,
Francesca Toccaceli and C alum McLennan were members of King's
College Cambridge at the time of the study (trustees, employees or
students).
CONFLICT OF INTEREST STATEMENT
The authors declare no conflict of interest.
DATA AVA ILAB ILITY STATE MEN T
Data are available from Dryad Digital Repository h t t p s: // do i .
org/10.5061/dryad.kd51c 5bbb (Marshall et al., 2023).
ORCID
Cicely A. M. Marshall https://orcid.org/0000-0002-7397-6472
Sebastian Eves- van den Akker https://orcid.
org/0000-0002-8833-9679
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SUPPORTING INFORMATION
Additional supporting information can be found online in the
Suppor ting Information section at the end of this article.
Table S1.Before-after-control-impact(BACI)hypothesistestingwith
ANOVA .
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Ecological Solutions and Evidence
MARSHALL et al.
Table S2. Emissions and costs associated with the mowing and
fertilising regimes of meadow and lawn in 2021, the two most
carbon intensive management activities.
Table S3. Species seed mix planted on the Back Lawn in October
20 19.
Figure S1. Pitfall biomass in 2021, showing the visual contrast
between meadow traps (top row) and lawn traps (bottom row).
Figure S2. Images of the meadow and lawn used to calculate relative
reflectance values.
Table S4. Relative reflectance values for analysed photographs.
Figure S3. Cambridge college greenspace, of which an estimated
56% (43.7 ha) is lawn.
Appendix S1. Survey questions.
How to cite this article: Marshall,C.A .M.,Wilkinson,M.T.,
Hadfield, P. M., Rogers, S. M., Shanklin, J. D., Eversham, B. C .,
Healey, R., Kranse, O. P., Preston, C. D., Coghill, S. J.,
McGonigle,K.L .,Moggridge,G.D.,Pilbeam,P.G.,Marza,A.
C.,Szigec san,D.,Mitchell,J.,Hicks,M.A.,Wallis,S.M.,Xu,
Z.…Eves-vandenAkker,S.(2023).Urbanwildflower
meadowplantingforbiodiversity,climateandsociety:An
evaluation at King's College, Cambridge. Ecological Solutions
and Evidence, 4, e12243. https://doi.org/10.1002/2688-
83 19.12 243
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