REVIEW ARTICLE
The impac o abandonmen and in ensi ica ion on he biodi e si y o
ag icul u ally ma ginal g asslands – a sys ema ic e iew
Susanna Hempel
a,b,*
, Felix He zog
a
, P´
e e Ba ´
a y
c,d
, E ik ¨
Ockinge
e
, E a Knop
a,b
a
Ag icul u al Landscapes and Biodi e si y, Ag oscope, Reckenholzs asse 191, Zu ich, Swi ze land
b
Depa men o E olu iona y Biology and En i onmen al S udies, Uni e si y o Zu ich, Win e hu e s asse 190, Zu ich, Swi ze land
c
Lendüle ’ Landscape and Conse a ion Ecology, Ins i u e o Ecology and Bo any, MTA–HUN-REN Cen e o Ecological Resea ch, V´
ac ´
a ´
o , Hunga y
d
Faunis ics and Wildli e Conse a ion, Depa men o Ag icul u e, Eco ophology, and Landscape De elopmen , Anhal Uni e si y o Applied Sciences, Be nbu g,
Ge many
e
Swedish Uni e si y o Ag icul u al Sciences, Depa men o Ecology, Uppsala, Sweden
ARTICLE INFO
Keywo ds:
Biodi e si y
Ex ensi e
In e eb a es
Plan s
T adi ional
Ve eb a es
ABSTRACT
Ag icul u ally ma ginal g asslands ha e been adi ionally managed a low in ensi y o cen u ies and a e among
Eu ope’s key biodi e si y ho spo s. Because o hei low p o i abili y, many o hem ha e been ei he abandoned
o subjec ed o in ensi ied managemen in ecen decades. Bo h pa hways h ea en he high di e si y o g assland
species ha depend on adi ional managemen p ac ices. To coun e ac he nega i e e ec s o abandonmen o
ag icul u al in ensi ica ion on biodi e si y, es o a ion and conse a ion p ac ices we e es ablished. Th ough a
sys ema ic li e a u e e iew based on 174 Eu opean s udies, we in es iga ed he impac s o abandonmen and
in ensi ica ion on a ious measu es o ag icul u ally ma ginal g asslands di e si y. Addi ionally, we ex ac ed
in o ma ion on he posi i e impac s o conse a ion – de ined he e as sus ained ex ensi e managemen p ac ices -
and es o a ion e o s aimed a p e iously abandoned o in ensi ied g asslands. Abandonmen had a high
p obabili y (71%) o educing plan and lichen biodi e si y, while i was signi ican ly less likely o dec ease he
di e si y o animals (23%). In ensi ica ion nega i ely a ec ed he di e si y o all o ganism g oups o a simila
ex en (65% p obabili y o plan s and lichen, 47% o animals). Conse a ion e o s we e likely o main ain o
inc ease animal biodi e si y (p obabili y 79%), bu in he s udies we analysed, hey we e no su icien o
p ese e he biodi e si y o plan s and lichen. The es o a ion o abandoned o in ensi ied g asslands was p e-
dic ed o enhance plan and lichen di e si y (68% p obabili y), while no signi ican ly changing animal di e si y.
Thus, di e en o ganisms g oups esponded di e en ly o changes in ag icul u al managemen , highligh ing he
need o a ge ed conse a ion and es o a ion s a egies. By syn hesizing biodi e si y esponses ac oss axa and
managemen ypes, his e iew con ibu es o a mo e in eg a ed and e idence-based unde s anding o how o
main ain and imp o e he ecological alue o ag icul u ally ma ginal g asslands.
In oduc ion
G asslands, one o he la ges e es ial biomes, co e a ound 40 %
o he Ea h’s e es ial su ace and domina e he landscape wo ldwide
(Gibson, 2023; Squi es e al., 2018). In addi ion o na u ally occu ing
g asslands, long pe iods o human ac i i y in a eas whe e na u al
ege a ion would o he wise no be he baceous ha e led o he de el-
opmen o semi-na u al g asslands (E iksson, 2020; Hejcman e al.,
2013). Cen u ies o adi ional managemen p ac ices, such as ex ensi e
mowing and g azing egimes, ha e shaped hese excep ionally
species- ich habi a s, which suppo a di e se a ay o plan and animal
li e (Dengle e al., 2014; Ellenbe g & Leuschne , 2010; Pe e mann &
Buzhdygan, 2021). Semi-na u al g asslands con ibu e o he conse a-
ion o many endemic and endange ed species (Ce abolini e al., 2016;
S oa e e al., 2009), and a e conside ed among he mos biodi e se
habi a s in Eu ope, ha bo ing mo e ascula plan species on a smalle
scale han opical ain o es s (Habel e al., 2013; Wilson e al., 2012).
Despi e hei s a us as biodi e si y ho spo s wi h high conse a ion
alue, only 4 % o semi-na u al g asslands a e p o ec ed wo ldwide,
making hem one o he mos h ea ened ecosys ems in he ace o cu -
en global change (P. T¨
o ¨
ok e al., 2016; Pe e mann & Buzhdygan,
2021).
* Co esponding au ho .
E-mail add ess: [email p o ec ed] (S. Hempel).
Con en s lis s a ailable a ScienceDi ec
Basic and Applied Ecology
jou nal homepage: www.else ie .com/loca e/baae
h ps://doi.o g/10.1016/j.baae.2025.08.003
Recei ed 11 Ap il 2025; Accep ed 11 Augus 2025
Basic and Applied Ecology 88 (2025) 9–18
A ailable online 12 Augus 2025
1439-1791/© 2025 The Au ho s. Published by Else ie GmbH on behal o Gesellscha ü Ökologie. This is an open access a icle unde he CC BY license (
h p://c ea i ecommons.o g/licenses/by/4.0/ ).
Ag icul u ally ma ginal g asslands a e a dis inc ype o low-
p oduc i e, semi-na u al g asslands, o en ound in a eas wi h shallow
soils, low nu ien a ailabili y, limi ed accessibili y o s eep slopes
(Richa ds e al., 2014). The economic sus ainabili y o managing such
ma ginal g asslands has sha ply declined in ecen decades, esul ing in
ei he ag icul u al in ensi ica ion o comple e abandonmen o land-use
p ac ices, wi h poo ly unde s ood consequences o biodi e si y (Ba ´
a y
e al., 2020; Shipley e al., 2024; S oa e e al., 2009). Gene ally, aban-
donmen , de ined as he cessa ion o adi ional managemen p ac ices,
leads o he na u al expansion o woody ege a ion. Ini ially, his can be
bene icial o biodi e si y by inc easing s uc u al complexi y, bu o e
ime i esul s in he loss o g assland-speci ic communi ies (Ma ini e al.,
2009; S. Fadda e al., 2008; Schmi & Rakosy, 2007). In con as ,
in ensi ica ion o ag icul u al p ac ices, such as he applica ion o
o ganic and mine al e ilize s combined wi h mo e equen mowing o
inc eased li es ock densi y, a o s as -g owing he baceous species,
leading o homogenized g assland communi ies domina ed by a ew
dis u bance- and nu ien - ole an species (Beckmann e al., 2019;
G een, 1990; Guo e al., 2023; Simons e al., 2015). This highligh s he
impo ance o low-in ensi y land-use p ac ices as a conse a ion s a egy
o p ese e he high biodi e si y o semi-na u al g asslands (Shipley
e al., 2024). On he o he hand, o es o e biodi e si y a e abandon-
men o in ensi ica ion, es o a ion p ojec s o en ein oduce adi-
ional mowing and g azing egimes o p e en succession by woody
species, o ein oduce los g assland di e si y by adding seeds o
plan ing na i e species, espec i ely (Lyons e al., 2023; Slodowicz e al.,
2023). Howe e , especially o ag icul u ally ma ginal g asslands, an
o e iew o he impac o abandonmen and in ensi ica ion on biodi-
e si y is lacking. Mos exis ing li e a u e consis s o indi idual s udies
ocused on local scales o speci ic species (bu see Ellio e al., 2023, o
a ecen e iew ega ding he impac o abandonmen ). Mo eo e ,
o e a ching e idence on he bene i s o biodi e si y managemen , such
as ag i-en i onmen schemes o na u e conse a ion p ac ices, emains
limi ed (Ba ´
a y e al., 2015).
We hus conduc ed a sys ema ic li e a u e e iew, agg ega ing in-
o ma ion om scien i ic s udies abou he consequences o manage-
men p ac ices on he biodi e si y o a ious g oups o species in
Eu opean ag icul u ally ma ginal g asslands. We ocused on he nega-
i e impac s o abandonmen and in ensi ica ion, while also add essing
he ex en o which conse a ion and es o a ion e o s can coun e ac
biodi e si y loss in hese species- ich semi-na u al ecosys ems. To ou
knowledge, his p o ides a new and in eg a ed pe spec i e ha com-
bines bo h h ea s and eco e y po en ial ac oss di e en o ganism
g oups in ag icul u ally ma ginal g asslands. This sys ema ic e iew
syn hesizes a ailable scien i ic in o ma ion o answe he ollowing
ques ions: 1) Wha a e he e ec s o abandonmen and in ensi ica ion on
he biodi e si y o ag icul u ally ma ginal g asslands, and do hese e -
ec s di e be ween lo al and aunal o ganism g oups. 2) Does he
ou come o abandonmen depend on he du a ion o abandonmen ? 3)
Does he ype o ag icul u ally ma ginal g assland in luence how o -
ganism g oups espond o in ensi ica ion o abandonmen ? 4) Do con-
se a ion and es o a ion e o s bene i biodi e si y ac oss di e en
o ganism g oups in ag icul u ally ma ginal g asslands ha aced aban-
donmen o in ensi ica ion? We hypo hesized ha bo h abandonmen
and in ensi ica ion educe he biodi e si y o ag icul u ally ma ginal
g asslands, wi h long- e m abandonmen ha ing mo e de imen al e -
ec s han sho - e m abandonmen . We hypo hesized ha lowe ophic
le els and less mobile o ganism g oups espond mo e s ongly and as e
o he nega i e e ec s o abandonmen o in ensi ica ion. Fu he mo e,
we hypo hesized ha conse a ion measu es help o main ain high
biodi e si y in hose habi a s, and ha es o a ion e o s mi iga e he
nega i e consequences o abandonmen and in ensi ica ion. By syn he-
sizing in o ma ion on he in luence o land-use p ac ices ac oss axo-
nomic g oups, his e iew con ibu es o a mo e in eg a ed
unde s anding o how biodi e si y esponds o wo con as ing an h o-
pogenic p essu es (in ensi ica ion and abandonmen ) on ag icul u ally
ma ginal g asslands. I also suppo s he g owing ield o e idence-based
conse a ion and es o a ion planning.
Me hods
Li e a u e sea ch and inclusion c i e ia
We conduc ed a li e a u e sea ch in Ap il 2023 using he Web o
Science Co e Collec ion (Science Ci a ion Index Expanded) and Else ie
Scopus da abases, employing a combina ion o keywo ds based on he
Popula ion, In e en ion, Compa ison, and Ou come (PICO) amewo k
(Higgins & G een, 2008). The PICO amewo k is a s uc u ed app oach
o sys ema ic e iews ha acili a es o mula ing esea ch ques ions by
de ining he Popula ion, In e en ion, Compa ison, and Ou come o
in e es , enabling a sys ema ic and comp ehensi e syn hesis o scien i ic
e idence. The ollowing de ini ions and inclusion c i e ia we e applied:
- Popula ion: We chose ag icul u ally ma ginal g asslands wi h
limi ed po en ial o in ensi e ag icul u al p oduc ion due o phys-
ical o abio ic cons ain s (e.g., oo d y, oo we , oo s eep, oo
emo e, o unsui able o be imp o ed by easonable amelio a ion
measu es) as he popula ion. The p ima y yield/ e u n o hese
ma ginal g asslands is g ass, hay o li e . This de ini ion includes
wooded meadows used o g azing o mowing, loodplain meadows,
and g azed ens.
- In e en ion: Rele an in e en ions included abandonmen ,
in ensi ica ion, o conse a ion o ex ensi ely used ma ginal g ass-
lands, as well as es o a ion o p e iously abandoned o in ensi ied
g asslands.
-Compa ison: Ma ginal g asslands ha we e main ained by ex en-
si e land-use p ac ices, such as long- e m adi ional mowing and
g azing egimes, we e chosen as con ol/compa a o .
- Ou come: The e ec s o ag icul u al managemen we e assessed
based on he ollowing biodi e si y me ics: species ichness, species
abundance, species di e si y, unc ional di e si y, o gene ic di-
e si y. Changes in biodi e si y alues we e ca ego ized as an in-
c ease, dec ease, o no change in esponse o ag icul u al
managemen .
The sea ch was es ic ed o s udy si es loca ed in Eu ope and
included only pee - e iewed a icles, con e ence p oceedings, and book
chap e s. We u he e ined he sea ch by limi ing i o speci ic Web o
Science ca ego ies and Scopus subjec a eas. To maximize he numbe o
po en ially ele an s udies, keywo ds we e linked acco ding o he
PICO amewo k using logical ope a o s. A de ailed s uc u e o he
PICO amewo k is p o ided in Appendix Table A1.
Exclusion c i e ia
The sea ch was conduc ed in Ap il 2023, yielding a o al o 3003
po en ially ele an s udies. We applied he P e e ed Repo ing I ems
o Sys ema ic Re iews and Me a-Analyses (PRISMA, Page e al., 2021)
checklis o guide he selec ion p ocess (Fig. 1).
A e emo ing duplica es, 1944 s udies emained and we e sc eened
in a h ee-s age p ocess ( i les, abs ac s, and ull- ex con en ) using he
ollowing exclusion c i e ia:
- No w i en in English o Ge man.
- Loca ed ou side he Eu opean con inen .
- The uni o s udy did no align wi h ou de ini ion o ag icul u ally
ma ginal g asslands (e.g., u ban g asslands, ield ma gins, bu ial
mounds, o cha ds, ineya ds, o o he ag o o es y sys ems).
- The in e en ion was un ela ed o local managemen changes (e.g.,
s udies ocusing on agmen a ion, habi a connec i i y, o landscape
complexi y).
- Re iews o modeling s udies.
S. Hempel e al.
Basic and Applied Ecology 88 (2025) 9–18
10
- Expe imen al s udies conduc ed in g eenhouses o labo a o ies.
Addi ionally, ou s udies ocusing on soil mic obes and ungi we e
excluded due o hei e y low ep esen a ion. A e applying hese
c i e ia, 174 s udies me he equi emen s ou lined in ou sys ema ic
e iew p o ocol.
Da a ex ac ion based on he consequences o g assland managemen on
di e en classes o o ganisms
In his sys ema ic e iew, we used he e m managemen synonymous
wi h land-use. We had p e-de ined wo majo ca ego ies o land-use
change occu ing in ag icul u ally ma ginal g asslands: abandonmen
and in ensi ica ion. Addi ionally, we ga he ed in o ma ion on conse -
a ion, which we de ined as he con inua ion o ex ensi e managemen
p ac ices, and es o a ion, which in ol es he ein oduc ion o adi-
ional managemen p ac ices o p e iously abandoned o in ensi ied
ag icul u ally ma ginal g asslands (Fig. 2).
Da a om he selec ed s udies we e eco ded in a s uc u ed da a
ex ac ion o m ocusing on managemen p ac ices (ca ego ized as
abandonmen , in ensi ica ion, conse a ion, o es o a ion), g assland
ypes (moun ain g asslands, calca eous and d y g asslands, empe a e
mesic and we g asslands), classes o o ganisms (plan s and lichen -
including ascula plan s, b yophy es and lichen; in e eb a es -
including insec s, a achnids, gas opods, and soil in e eb a es; and
e eb a es - such as bi ds and small mammals), and changes in a ious
biodi e si y alues (species ichness, species abundance, species di-
e si y, unc ional di e si y, o gene ic di e si y). These changes we e
ca ego ized as dec ease (−1), no change (0), o inc ease (1) based on
s a is ical signi icance and con idence in e als p esen ed in ex , ig-
u es, o ables (Appendix B). Desc ip i e isualisa ions o he aw da a
can be ound in Appendix Fig. A1 ep esen ing he mean esponse o
single o de s o o ganisms o managemen egimes, and Appendix
Fig. A2 showing he mean esponse o single biodi e si y measu es o
land-use change.
To assess he e ec s o ongoing succession on biodi e si y, we
u he ca ego ized s udy si es in o h ee classes based on he ime since
abandonmen : Sho - e m abandonmen (<5 yea s) was classi ied as
ea ly s ages o succession, whe e g assland species emain dominan and
s uc u al changes a e minimal. Mid- e m abandonmen was de ined as
an abandonmen o 5–10 yea s. Fo his ime ange o abandonmen
¨
Ockinge e al. (2006) ha e ound a beginning o sh ub enc oachmen
and he es ablishmen o compe i i e g asses, bu a pe sis ence o mos
semi-na u al g assland species . Long- e m abandonmen was de ined as
>10 yea s a e abandonmen and is expec ed o co espond o s uc-
u al and composi ional changes, po en ially accompanied by he
es ablishmen o woody ege a ion and he educ ion o open habi a .
Each a icle epo ed one o mo e managemen - ela ed changes in
biodi e si y, enabling us o ex ac a o al o 440 eco ded obse a ions
ega ding he consequences o land-use p ac ices on di e en classes o
o ganisms ac oss he 174 s udies. We applied s ic selec ion c i e ia and
igo ously e alua ed he epo ed impac s o ag icul u al managemen
on biodi e si y alues. The qual ia i e syn hesis allowed us o include a
la ge numbe o s udies and in eg a e highly he e ogeneous da ase s,
ensu ing a b oad ep esen a ion o he e idence base in he ield and
p o iding a comp ehensi e analysis o ou esea ch ques ion.
Da a analysis
In he desc ip i e analysis, we ca ego ized o ganisms in o h ee
g oups: plan s and lichens, in e eb a es, and e eb a es. Howe e , o
he s a is ical analysis, we combined he in e eb a e and e eb a e
ca ego ies in o a single g oup, called animals, due o he limi ed numbe
o s udies ocused on e eb a es.
To assess he nega i e e ec s o abandonmen o in ensi ica ion on
he biodi e si y o ag icul u ally ma ginal g asslands, we assigned a
alue o 1 o each ins ance whe e a dec ease in biodi e si y was
obse ed, and a alue o 0 o ins ances whe e biodi e si y ei he
inc eased o emained unchanged.To e alua e he posi i e e ec s o
conse a ion on biodi e si y, we assigned a alue o 1 o obse a ions
whe e biodi e si y ei he did no change o inc eased (i.e., whe e con-
se a ion ac ions p e en ed biodi e si y loss), and a alue o 0 o
Fig. 1. PRISMA (P e e ed Repo ing I ems o Sys ema ic Re iews and Me a-Analyses) low diag am epo ing he numbe o s udies iden i ied and excluded du ing
he sc eening p ocess.
Fig. 2. G aphic ep esen a ion o land-use p ac ices in ag icul u ally ma ginal
g asslands ha we e conside ed in his sys ema ic e iew. Blue numbe s indi-
ca e he amoun o obse a ions ega ding he consequences on biodi e si y o
each change in managemen p ac ice o in ensi y.
S. Hempel e al.
Basic and Applied Ecology 88 (2025) 9–18
11
obse a ions epo ing a dec ease in biodi e si y.Finally, o analyze he
posi i e e ec s o es o a ion on abandoned o in ensi ied g asslands, we
assigned a alue o 1 o obse a ions epo ing an inc ease in biodi-
e si y and a alue o 0 o hose indica ing no change o a dec ease in
biodi e si y (Table 1).
Fo analyzing he nega i e impac o abandonmen and in ensi ica-
ion o ag icul u ally ma ginal g asslands on he di e si y o di e en
g oups o o ganisms (ques ion 1), we calcula ed he p obabili y o
obse ing a signi ican decline in biodi e si y ollowing abandonmen o
in ensi ica ion. We i ed a gene alized linea mixed-e ec s model wi h a
binomial dis ibu ion, using change in biodi e si y as he esponse
a iable, he in e ac ion be ween managemen (abandonmen , in ensi-
ica ion) and o ganism g oup (plan s and lichen, animals) as p edic o s,
and s udy ID as a andom e ec o accoun o a iabili y be ween
s udies.
The limi ed numbe o obse a ions o each ype in ensi ica ion
p ecluded a o mal s a is ical analyses. Ne e heless, o a desc ip i e
analysis, we calcula ed he mean bina y esponse o obse ing a
dec ease in biodi e si y upon he di e en in ensi ica ion measu es.
To answe ques ion 2, ega ding whe he he nega i e impac o
abandonmen depends on i s du a ion, we calcula ed he p obabili y o
obse ing a signi ican decline in biodi e si y ollowing di e en du a-
ions o abandonmen . We i ed a gene alized linea mixed-e ec s
model wi h a binomial dis ibu ion, using change in biodi e si y as
he esponse a iable, he in e ac ion be ween o ganism g oup (plan s
and lichen, animals) and du a ion o abandonmen (long- e m, mid-
e m, sho - e m) as p edic o s, and s udy ID as a andom e ec .
To de e mine whe he di e en ypes o ag icul u ally ma ginal
g asslands espond di e en ly o abandonmen o in ensi ica ion
(ques ion 3), we calcula ed he p obabili y o obse ing a signi ican
dec ease in biodi e si y by i ing a gene alized linea mixed-e ec s
model wi h a binomial dis ibu ion using change in biodi e si y as he
esponse a iable, he in e ac ion be ween managemen (abandonmen ,
in ensi ica ion), o ganism g oup (plan s and lichen, animals) and
g assland ype (moun ain g asslands, d y and calca eous g asslands,
empe a e mesic and we g asslands) as p edic o s, and s udy ID as a
andom e ec .
Fo ques ion 4 conce ning he bene icial e ec s o conse a ion and
es o a ion on p ese ing he di e si y o di e en g oups o o ganisms
(plan s and lichen, animals), we calcula ed he p obabili y o obse ing
an inc ease in biodi e si y a e es o a ion and he p obabili y o no
change o an inc ease in biodi e si y ollowing conse a ion e o s, by
i ing a gene alized linea mixed-e ec s model wi h a binomial dis i-
bu ion, using change in biodi e si y as he esponse a iable, he
in e ac ion be ween managemen (conse a ion, es o a ion) and o -
ganism g oup (plan s and lichen, animals) as p edic o s, and s udy ID as
a andom e ec .
To assess he signi icance o indi idual ca ego ies wi hin he model
e ms, we e i ed he model wi hou an in e cep . This app oach es i-
ma es absolu e e ec s o each ca ego y a he han ela i e di e ences
compa ed o a e e ence le el, allowing di ec e alua ion o he signi -
icance o each ca ego y (Ag es i, 2007). We e alua ed he signi icance o
indi idual ca ego ies and di e ences o he e e ence le el using wo
complemen a y app oaches. Fi s , we examined he p- alues om he
model summa y ou pu o assess whe he indi idual p edic o s
signi ican ly in luenced he esponse a iable. Second, we isualized
p edic ed p obabili ies wi h 95 % con idence in e als (CIs) o in e p e
he esul s. As we used binomial models, we conside ed an e ec sig-
ni ican i he con idence in e al did no include 0.5, indica ing a clea
p obabili y shi . Addi ionally, o compa isons be ween ca ego ies, we
assessed signi icance based on whe he he con idence in e als o e -
lapped, whe e subs an ial o e lap sugges ed no clea di e ence be ween
g oups.
The s a is ical analyses we e pe o med using he idy e se (Wickham
e al., 2019) and glmmTMB packages (B ooks e al., 2017) in he ee-
wa e R s a is ical so wa e ( e sion R 4.4.2, h p:// -p ojec .o g).
Resul s
Geog aphical and empo al dis ibu ion o scien i ic e idence
We iden i ied 174 scien i ic s udies add essing managemen p ac-
ices in ag icul u ally ma ginal a eas ac oss Eu ope, om which we
ex ac ed 440 obse a ions: 230 deal wi h abandonmen , 110 wi h
in ensi ica ion, 40 wi h conse a ion, and 60 wi h es o a ion (Fig. 2).
Among hese 174 s udies, he numbe o ele an publica ions has
inc eased o e he pas h ee decades (Appendix Fig. A3), sugges ing
g owing ecogni ion o his issue.
The s udies included in he inal analysis encompassed nea ly all
egions o he Eu opean con inen , apa om he a Eas e n a eas
(Fig. 3). The dis ibu ion o obse a ions was ela i ely simila o all
ypes o managemen , wi h mos o he s udies being conduc ed in
Cen al Eu opean coun ies.
E ec o di e en managemen p ac ices on biodi e si y o ag icul u ally
ma ginal g asslands
The numbe o obse a ions abou he e ec s o ag icul u al man-
agemen on he di e si y o ma ginal g asslands (Fig. 4) was highes o
plan s (211 obse a ions) and in e eb a es (206 obse a ions), wi h
conside ably ewe s udies ocusing on e eb a e biodi e si y (23 ob-
se a ions). The e was a no iceable bias owa ds ascula plan s (210
obse a ions) and p ominen insec g oups such as bu e lies (55 ob-
se a ions), bee les (40 obse a ions) and g asshoppe s (29 obse a-
ions). Only a ew s udies add essed he e ec s o land-use p ac ices in
ag icul u ally ma ginal g asslands on he di e si y o b yophy es (7
obse a ions), lichens (8 obse a ions), gas opods (11 obse a ions),
an s (8 obse a ions), and spide s (19 obse a ions). Addi ionally, jus
one s udy in es iga ed he impac o g assland managemen on small
mammal di e si y (To e & Palau, 2023). The limi ed numbe o s udies
examining he e ec o land-use p ac ices on p o is s and ungal di-
e si y p ecluded hei inclusion in ou analysis.
Due o he limi ed numbe o obse a ions ocusing on he conse-
quences o g assland managemen on e eb a es, we combined he
in e eb a e and e eb a e ca ego ies o he s a is ical analysis in o a
single g oup (animals).
In line wi h ou i s ques ion, we ound a signi ican ly high p oba-
bili y o obse ing a decline in di e si y o plan s and lichens due o
abandonmen (71 % [59 % o 81 %], Fig. 5, Appendix Table A2).
Con e sely, animals showed a signi ican ly low p obabili y o di e si y
decline ollowing abandonmen (23 % [15 % o 34 %]), which was also
signi ican ly lowe han o plan s (Fig. 5, Appendix Table A2).
Rega ding ou second ques ion we ound a signi ican ly high p ob-
abili y o obse ing a decline in di e si y o plan s and lichen a e long-
e m abandonmen o mo e han 10 yea s (76 % [60 % o 87 %], Ap-
pendix Fig. A4, Appendix Table A3). While he p edic ed p obabili ies
o mid- e m (5 – 10 yea s) and sho - e m (<5 yea s) abandonmen
we e simila in magni ude, hei con idence in e als included 0.5 and
we e he e o e no s a is ically signi ican (Appendix Fig. A4, Appendix
Table A3). The p obabili y o obse ing a decline in animal di e si y in
long- e m abandoned plo s was 39 % [26 % o 53 %], which was
Table 1
O e iew o managemen p ac ices, and hei e ec on biodi e si y wi h co -
esponding dis ibu ion o he binomial alues o s a is ical modeling.
Managemen
p ac ice
E ec on
biodi e si y
Dec ease No
change
Inc ease
Abandonmen nega i e 1 0 0
In ensi ica ion nega i e 1 0 0
Conse a ion posi i e/no e ec 0 1 1
Res o a ion posi i e 0 0 1
S. Hempel e al.
Basic and Applied Ecology 88 (2025) 9–18
12
signi ican ly highe compa ed o sho - e m (8 % [2 % o 28 %]) o mid-
e m abandonmen (7 % [2 % o 25 %]), bo h o which we e signi ican ly
low (Appendix Fig. A4, Appendix Table A3).
Fo ou hi d ques ion, which examined he e ec o abandonmen o
in ensi ica ion on he biodi e si y o di e en g assland ypes, we ound
a signi ican ly high p obabili y o obse ing a decline in plan and lichen
di e si y a e abandonmen o moun ain g asslands (81 % [62 % o 92
%], Appendix Fig. A5, Appendix Table A4). The p obabili ies o a decline
in plan and lichen di e si y a e abandonmen we e 67 % [46 % o 83
%] in d y and calca eous g asslands, and 62 % [39 % o 80 %] in
empe a e mesic and we g asslands (Appendix Fig. A5, Appendix
Table A4). Fu he mo e, he e was a signi ican ly low p obabili y o
obse ing a decline in animal di e si y a e abandonmen o d y and
calce ous (20 % [9 % o 39 %]), and empe a e mesic and we g asslands
(20 % [10 % o 38 %], Appendix Fig. A5, Appendix Table A4). The
p obabili y o de ec ing a decline in animal di e si y a e abandonmen
in moun ain g asslands was 31 % [15 % o 53 %] (Appendix Fig. A5,
Appendix Table A4).
Fu he analysing ou i s and hi d ques ion, he p obabili y o
obse ing a decline in biodi e si y a e in ensi ica ion was simila ly
Fig. 3. Geog aphical dis ibu ion o obse a ions om s udies included in he inal analysis. Ci cle size is scaled o isualiza ion pu poses o imp o e isibili y o
o e lapping da a poin s, and does no encode any quan i i e in o ma ion. The cen e o each ci cle co esponds o he coo dina es o he eseach a ea o he
included s udies.
Fig. 4. O e iew o obse a ions epo ing an e ec o managemen on he di e si y o di e en classes o o ganisms in ag icul u ally ma ginal g assland.
S. Hempel e al.
Basic and Applied Ecology 88 (2025) 9–18
13
high o bo h, plan s and lichen as well as animals (65 % [46 % o 80 %]
and 47 % [32 % o 63 %], espec i ely, Fig. 5, Appendix Table A2), and
did no di e be ween di e en g assland ypes (Appendix Fig. A5,
Appendix Table A4). The in ensi ica ion measu es analyzed in he
e iewed s udies p ima ily included in ensi e g azing o mowing, ei he
sepa a ely o in combina ion wi h e iliza ion. Inc eased e iliza ion
was p edominan ly applied alongside mowing, a he han as an isola ed
measu e o in combina ion wi h g azing. Due o limi ed sample sizes, a
o mal s a is ical analysis was no easible, and compa isons we e
ins ead based on he p opo ion o cases in which in ensi ica ion led o a
dec ease in biodi e si y (mean bina y esponse, Appendix Fig. A6).This
gene al assessmen o pa e ns sugges ed ha plan s a e mo e suscep-
ible o he combined e ec s o in ensi e mowing and e iliza ion
compa ed o animals, whe eas in ensi e mowing alone had a simila
impac on bo h g oups o o ganisms (Appendix Fig. A6). Addi ionally,
in ensi e i iga ion appea ed o nega i ely a ec plan di e si y, bu no
animal di e si y (Appendix Fig. A6).
Ad essing ou ou h ques ion, he p obabili y o main aining o
inc easing he biodi e si y o animals was signi ican ly high (79 % [55
% o 92 %]) a e conse a ion o ex ensi ely managed ag icul u ally
ma ginal g asslands, whe eas i showed a signi ican ly lowe p obabili y
(41 % [20 % o 66 %]) o main aining o imp o ing he di e si y o
plan s and lichen (Fig. 6, Appendix Table A5). The signi icance o he
di e ence in conse a ion e ec s be ween plan s and lichen s. animal
di e si y was assessed using he model summa y ou pu (Appendix
Table A5), whe e he p- alue o he con as is 0.027. Fu he mo e, he
con idence in e als do no subs an ially o e lap, suppo ing a signi i-
can ly s onge e ec o conse a ion on animals han on plan s and
lichens.
The p obabili y o obse ing an inc ease in di e si y o plan s and
lichen was signi ican ly high a e es o a ion o p e iously abandoned
o in ensi ied ag icul u ally ma ginal g asslands (68 % [52 % o 80 %],
Fig. 6, Appendix Table A5). In con as , he p obabili y o obse ing an
inc ease in animal di e si y upon es o a ion – which, acco ding o ou
model, did no signi ican ly decline ollowing abandonmen o in ensi-
ica ion (Fig. 5) – was 44 % [20 % o 70 %] (Fig. 6, Appendix Table A5).
Discussion
This sys ema ic li e a u e e iew is comp ised o 174 s udies con-
duc ed in Eu ope o e he las 30 yea s and ocused on he e ec s o
abandonmen and in ensi ica ion on biodi e si y o ag icul u ally ma -
ginal g asslands, while also assessing he ole o conse a ion and
es o a ion. We combined and classi ied an inhomogeneous co pus o
s udies in o s a is ical models, which allowed us o e eal con as ing
e ec s o managemen changes. Ou indings la gly con i m ou hy-
po heses: bo h abandonmen and in ensi ica ion end o educe biodi-
e si y, wi h long- e m abandonmen ha ing a mo e de imen al e ec
han sho - o mid- e m abandonmen . Lowe ophic le els and less
mobile o ganism g oups, pa icula ly plan s and lichens, we e mo e
Fig. 5. P obabili ies o obse ing a dec ease in biodi e si y upon abandonmen o in ensi ica ion o ag icul u ally ma ginal g asslands. Numbe s nex o da apoin s
ep esen sample sizes. Raw da a a e o e laid as ji e ed poin s o show he dis ibu ion o obse ed ou comes ha in o med he model.
Fig. 6. P obabili ies o obse ing an inc ease o no change in biodi e si y upon conse a ion o ex ensi ely managed g asslands, and o obse ing an inc ease in
biodi e si y a e es o a ion o p e iously abandoned o in ensi ely used ag icul u ally ma ginal g asslands. Numbe s indica e sample sizes. Raw da a a e o e laid as
ji e ed poin s o show he dis ibu ion o obse ed ou comes ha in o med he model.
S. Hempel e al.
Basic and Applied Ecology 88 (2025) 9–18
14
s ongly and apidly a ec ed by abandonmen han animals. Fu he -
mo e, ou esul s suppo he hypo hesis ha conse a ion ac ions help
main ain high biodi e si y, mos no ably o animals, while es o a ion
e o s seem pa icula ly e ec i e in mi iga ing biodi e si y loss among
plan communi ies.
E ec s o abandomen
This sys ema ic e iew con i ms he dec ease in plan di e si y due
o abandonmen and demons a es ha moun ain g asslands a e
pa icula ly sensi i e o he lack o dis u bance compa ed o o he
ag icul u ally ma ginal g asslands. The cessa ion o low-in ensi y man-
agemen p ac ices in ag icul u ally ma ginal g asslands leads o an
accumula ion o biomass, which gene ally has nega i e consequences
o plan communi ies (Dengle e al., 2014; Peco e al., 2017; Valko
e al., 2018). One eason o he decline in plan di e si y due o man-
agemen abandonmen migh be ha plan li e al e compe i ion o
ligh a ailabili y, a o ing less ligh -demanding species, ul ima ely
homogenising g assland species composi ion (J. Valk´
o e al., 2018).
Thus, ou indings s ess he impo ance o low-in ensi y managemen
(e.g., biomass emo al by mowing, g azing, bu ning) i we we e o
conse e he high plan di e si y o ag icul u ally ma ginal g asslands.
The compa a i ely weake esponse o d y and calca eous, as well as
empe a e mesic and we g asslands o abandonmen may e lec di -
e ences in hei ini ial plan communi y composi ion, land-use his o y,
and en i onmen al condi ions. Fo example, Pop che a e al. (2009)
ound no clea co ela ion be ween ime since abandonmen and plan
species ichness in we g asslands, which hey a ibu ed o si e-speci ic
di e ences in successional ajec o ies. G asslands domina ed by
s ess- ole an species and occu ing on low-nu ien soils may expe i-
ence slowe ege a ion shi s and less p onounced biodi e si y loss a e
abandonmen , while hose wi h nu ien -demanding o
dis u bance-adap ed species may change mo e apidly. Mo eo e , he
unc ional ai s and compe i i e s a egies o dominan species likely
media e how plan communi ies espond o he cessa ion o manage-
men (Pop che a e al., 2009). This ecological he e ogenei y may
explain he mo e mode a e biodi e si y esponses we obse ed in
empe a e mesic and we , as well as d y and calca eous g assland ypes
compa ed o moun ain g asslands.
In con as o he esponse o plan s and lichen, we ound ha
abandonmen o ag icul u ally ma ginal g asslands had a signi ican ly
lowe p obabili y o dec easing animal di e si y, con i ming he ind-
ings o Ellio e al. (2023). This e ec can likely be a ibu ed o se e al
ac o s. Fi s ly, he g ea di e si y o axonomic g oups wi hin in-
e eb a es and e eb a es may esul in di e se esponses o land-use
p ac ices, in luenced by ac o s such as mobili y, habi a ange,
eeding, and nes ing beha io s and hus a sys ema ic esponse on he
le el o animals is no de ec able (E ns e al., 2017). Fo example, some
species may be mo e mobile and able o mig a e o sui able habi a s in
esponse o managemen changes. In con as , species wi h limi ed
mobili y may be mo e ulne able o habi a deg ada ion (L¨
o le e al.,
2020). Addi ionally, di e ences in eeding habi s, such as he bi o ous
e sus ca ni o ous die s, may cause a ying deg ees o dependence on
plan communi ies o p ey a ailabili y, u he in luencing species’ e-
sponses o changes in he landscape (S. Fadda e al., 2008). Secondly,
highe ophic le els (e.g., he bi o es and p eda o s) may exhibi a
delayed esponse o changes in lowe ophic le els, such as plan s.
Thi dly, he abandonmen o g assland managemen ini ially inc eases
s uc u al complexi y, po en ially bene i ing ce ain a h opod and bi d
popula ions (Azca a e & Peco, 2012; Koch e al., 2015; Laiolo e al.,
2004; P¨
oy y e al., 2006). Howe e , long- e m abandonmen may cause
a decline in g assland-speci ic in e eb a e species due o educed
habi a he e ogenei y a e bush enc oachemen , subsequen ly a ec ing
insec i o ous bi ds (Bau e al., 2006; O. Valk´
o e al., 2018). We ocused
ou analysis p ima ily on he impac s o managemen p ac ices on
biodi e si y in ag icul u ally ma ginal g asslands by examining
indica o s ela ed o species ichness, abundance, species di e si y,
unc ional di e si y and gene ic di e si y. Ye , e en when hose di-
e si y indica o s emain simila , al e a ions in species iden i y and
he e o e communi y composi ion may occu .
E ec s o in ensi ica ion
Al hough ag icul u ally mo e p oduc i e and accessible ields ha e
his o ically aced g ea e p essu e om in ensi ica ion (Emme son e al.,
2016), ou e iew s ill iden i ied 110 s udies in es iga ing in ensi ica-
ion in ag icul u ally ma ginal g asslands, which a e ecosys ems ypi-
cally less sui ed o high-in ensi y use due o opog aphical and
en i onmen al cons ain s (Meie e al., 2022). Ou analysis shows ha
e en modes le els o in ensi ica ion nega i ely a ec ed he di e si y o
plan s and lichen as well as animals o a simila ex end - 65 % and 47 %
p obabili y, espec i ely. These p obabili ies o biodi e si y decline
ollowing in ensi ica ion did no each s a is ical signi icance, which
may e lec he compa a i ely lowe in ensi y o p ac ices applied in
hese a eas. The simila esponse ac oss axonomic g oups is consis en
wi h nume ous s udies showing ha e en mode a e in ensi ica ion can
lead o apid habi a homogeniza ion, a o he dominance o
as -g owing and dis u bance- ole an species, and ou compe e na i e
g assland specialis s. This no only al e s communi y composi ion bu
also dis up s bio ic in e ac ion ne wo ks, as well as weakens he pai -
wise di e si y co ela ions among axa, wi h s ong land-use in ensi i-
ca ion e en b eaking specialized species ela ionships, ul ima ely
con ibu ing o widesp ead biodi e si y loss in ma ginal g asslands
(Beckmann e al., 2019; G een, 1990; Guo e al., 2023; Manning e al.,
2015; McKeon e al., 2022). On a la ge spa ial scale, he decline in
bio ic he e ogenei y due o land-use in ensi ica ion esul s in educed
composi ional dissimila i y among di e en si es, which poses a majo
h ea o o e all biodi e si y (Gossne e al., 2016).
Fu he mo e, in ensi ica ion was p edic ed o ha e a signi ican ly
mo e de imen al e ec on animal di e si y han abandonmen , po en-
ially because he consequences o in ensi ied managemen p ac ices
impac he di e si y o g assland species conside ably as e han he
slow succession occu ing a e land abandonmen (L¨
o le e al., 2020).
E ec s o conse a ion measu es and es o a ion e o s
The scien i ic s udies ha we iden i ied in his sys ema ic e iew
ocussed on he impac o abandonmen o in ensi ica ion on he di-
e si y o ma ginal g asslands. Fi y o hese s udies also in es iga ed he
e ec s o conse a ion o es o a ion p ac ices. As ou sea ch s ing did
no include speci ic e ms ela ed o conse a ion o es o a ion, we do
no claim o p o ide an exhaus i e o e iew o hei e ec s. None he-
less, gi en hei c i ical impo ance in shaping biodi e si y ou comes in
ag icul u ally ma ginal g asslands, we sough o inco po a e ele an
in o ma ion whe e a ailable, aiming o p o ide a b oade con ex and
highligh he ole o conse a ion and es o a ion e o s alongside he
impac s o abandonmen and in ensi ica ion.
The g assland conse a ion e o s co e ed in he s udies included in
ou analysis in ol ed adi ional meadow i iga ion (Schi mel & Ge -
lach, 2022), bu ning (Dmy ash-Va seba & Shumska, 2020; Ham ik &
Kosulic, 2021; K¨
ohle e al., 2005), mechanical u dis u bance (Ham ik
& Kosulic, 2021), mulching (Cabon e al., 2021; Dolezal e al., 2011;
Gaisle e al., 2019; Pe ez-Sanchez e al., 2018), al e na i e
low-in ensi y mowing egimes (Kenye es & Szen i mai, 2017), g azing
wi h low s ocking a e (Bona i e al., 2017; P. T¨
o ¨
ok e al., 2016; and
o he s), g azing on sowing en iched pas u es (Mo eno-Opo e al., 2021),
and lea ing uncu e uges (Kalab e al., 2020; R´
e ´
esz e al., 2025). Ou
esul s show ha hese conse a ion e o s a e p edic ed o main ain o
e en inc ease he di e si y o in e eb a es and e eb a es, al hough
hey a e no p edic ed o be as e ec i e in main aining o inc easing he
di e si y o plan s and lichen. This aises conce ns ha low-in ensi e
conse a ion ac ions migh no always be su icien o main ain he
S. Hempel e al.
Basic and Applied Ecology 88 (2025) 9–18
15
high lo al di e si y o p e iously ex ensi ely g azed o mown ag icul-
u ally ma ginal g asslands in he ace o clima e change and species
in asion.
Res o a ion e o s, on he o he hand, we e associa ed wi h a high
p edic ed p obabili y o posi i ely a ec ing he di e si y o plan s and
lichen, which also showed he highes likelihood o decline unde
abandonmen , bu we e also nega i ely impac ed by in ensi ica ion ac-
co ding o ou analysis. Howe e , he di e si y o animals was no
p edic ed o signi ican ly inc ease as a esul s o es o a ion e o s,
which is expec ed, gi en ha hese axa had a low p edic ed p obabili y
o being nega i ely a ec ed by abandonmen and a mode a e p oba-
bili y o being impac ed by in ensi ica ion in he i s place. The es o-
a ion o abandoned o in ensi ied ma ginal g asslands in he s udies we
analyzed was p edominan ly achie ed h ough he ein oduc ion o
ex ensi e g azing o mowing (Colom e al., 2021; Rysiak e al., 2021;
and o he s). Abandoned g asslands we e u he es o ed by sh ub
clea ing (Baba, 2003; Campedelli e al., 2016; ¨
Ockinge e al., 2006;
Zeidle e al., 2022). These e o s seem su icien o inc ease plan di-
e si y ha su e ed om he consequences o abandonmen o in en-
si ica ion. Fu he mo e, a ecen s udy demons a ed ha es o a ion
h ough plan ma e ial ans e p omo ed endange ed species (Somme
e al., 2025).
In o de o comba global g assland deg eda ion an in eg a ed socio-
ecological app oach is equi ed ha in ol es inc easing he ecogni ion
o g asslands in policy amewo ks, de eloping s anda dized indica o s
o assessing deg eda ion and es o a ion, employing scien i ic knowl-
edge sha ing and s akeholde engagemen , o add ess he d i e s o
deg ada ion, acili a e a ge ed es o a ion e o s, and ensu e ha
ecological and socio-economic bene i s a e p ese ed and enhanced
(Ba dge e al., 2021).
Limi a ions and di ec ions o u u e esea ch
While ou sys ema ic e iew o e s impo an insigh s in o how
di e en managemen egimes a ec biodi e si y in ag icul u ally
ma ginal g asslands, we we e cons ained o g ouping o ganism e-
sponses in o b oade ca ego ies – “plan s and lichens” s. “animals” –
due o insu icien da a o many speci ic axa. This p e en ed mo e ine-
scale analyses ac oss axonomic g oups and may mask impo an axon-
speci ic pa e ns. Fu u e esea ch should aim o ill hese gaps by
inc easing he axonomic and ecological esolu ion o biodi e si y
moni o ing in g assland sys ems. In he long e m, his will enable mo e
de ailed syn heses, ideally in he o m o o mal me a-analyses, which
would allow o obus e ec size es ima ion and be e assessmen o
a iabili y among axa, habi a ypes, and managemen p ac ices.
Conclusions
In conclusion, ou sys ema ic e iew highligh s he di e en ial im-
pac s o managemen changes on biodi e si y o ag icul u ally ma ginal
g asslands. While abandonmen poses a subs an ial h ea o plan and
lichen di e si y, i s nega i e e ec on animal di e si y is conside ably
less p onounced. Ag icul u al in ensi ica ion simila ly a ec s biodi e -
si y ac oss he in es iga ed o ganism g oups. Encou agingly, conse a-
ion ac ions – de ined he e as he con inua ion o adi ional, low-
in ensi y managemen – a e likely o main ain o enhance animal di-
e si y, hough hey we e no consis en ly e ec i e o plan s and li-
chens. Res o a ion e o s, in u n, we e mos p omising o plan s and
lichens.
Ul ima ely, ou indings emphasize ha no single managemen
s a egy bene i s all axa equally and highligh he need o axon-
speci ic and con ex -sensi i e managemen s a egies ha e lec he
a ying ecological needs o di e en o ganism g oups. Gi en ha bo h
abandonmen and in ensi ica ion unde mine biodi e si y, albei in
axon-speci ic ways, policy amewo ks should p e en ei he in ag i-
cul u ally ma ginal g asslands. This is especially impo an in egions
whe e abandonmen leads o apid successional o e g ow h and whe e
in ensi ied use exceeds he ecological ca ying capaci y o he habi a .
Ag i-en i onmen schemes should he e o e be e ined o be e e lec
he land-use his o y, g assland ype and a ge axa o each si e, e.g.,
di e en ia ing be ween d y calca eous g asslands and we meadows in
subsidy design. Finally, in eg a ing long- e m moni o ing in o conse -
a ion and es o a ion p og ams will be essen ial o ack how di e en
o ganism g oups espond o in e en ions and o suppo adap i e,
e idence-based g assland managemen .
Funding
This s udy has ecei ed unding om he Eu opean Union’s Ho izon
2020 esea ch and inno a ion p og amme unde g an ag eemen No
862,480 ( he SHOWCASE p ojec ) and om he Fede al O ices o
Ag icul u e and o he En i onmen ( he ALL-EMA p ojec ).
CRediT au ho ship con ibu ion s a emen
Susanna Hempel: W i ing – e iew & edi ing, W i ing – o iginal
d a , Me hodology, In es iga ion, Fo mal analysis, Da a cu a ion,
Concep ualiza ion. Felix He zog: W i ing – e iew & edi ing, Supe i-
sion, Funding acquisi ion, Concep ualiza ion. P´
e e Ba ´
a y: W i ing –
e iew & edi ing, Concep ualiza ion. E ik ¨
Ockinge : W i ing – e iew &
edi ing, Concep ualiza ion. E a Knop: W i ing – e iew & edi ing, Su-
pe ision, Me hodology, Funding acquisi ion, Concep ualiza ion.
Decla a ion o compe ing in e es
The au ho s decla e ha hey ha e no known compe ing inancial
in e es s o pe sonal ela ionships ha could ha e appea ed o in luence
he wo k epo ed in his pape .
Supplemen a y ma e ials
Supplemen a y ma e ial associa ed wi h his a icle can be ound, in
he online e sion, a doi:10.1016/j.baae.2025.08.003.
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