View metadata,citationandsimilarpapersatcore.ac.uk This article isThis article by protected copyright. All rightsreserved. 10.1111/ibi.12696 doi: differences to lead between the of thisversion c and Version Record.Please copyediting, paginationbeen throughthe andproofreadingtypesetting, process,may which This article acceptedhas been for publication andundergone fullpeer review buthasnot one treatments A recently deepor threeyears over examined responses ground whether To determine E * Running Ma Zhijun Editor: Handling Short Communication Article Type: Experimental evidence ROBERT W. HAWKES, W. ROBERT mail bundance increased through the study, the study, through increased bundance C

Accepted author: orresponding Article -

or two or :

[email protected] 4 - 2 - Natural England, England, Natural cultivated) and complexity (homogenous or ‘complex or (homogenous andcomplexity cultivated) RSPB Centre for Conservation Science, The Lodge, Sandy, Bedfordshire, SG19 2DL, UK 2DL, SG19 Bedfordshire, Sandy, The Science, Lodge, Conservation for Centre RSPB head cultivated subplots), plus controls, replicated across the U the replicated across controls, plus subplots), cultivated 1 School of Environmental Sciences, University of East Anglia, Norwich, NR4 7TJ, UK 7TJ, NR4 Norwich, Anglia, East UniversitySciences, of Environmental School of . Within complex . - 3 year School of Biological Sciences, University of East Angl ofEast University Sciences, Biological School of : responses to ground responses Woodlark - old fallows. Regardless of of Regardless fallows. old 1 * JENNIFER. SMART, JENNIFER. *

Suite D Unex House Bourges Boulevard, Peterborough, PE11NG Peterborough, Boulevard, Unex Bourges House D Suite

- mosaics - disturbance increased Woodlark Woodlark increased disturbance

that ,

to four four to Woodlark preferentially used recently used preferentially Woodlark Lullula arborea and was and was

ground 2 treatment , 3 treatments treatments

- ANDY. BROWN, ANDY. disturbance higher on plots close plots higher on

-

disturbance benefits Woodlark detail, providing suitable foraging habitat habitat foraging suitable providing detail, varying in establishment method (shallow method inestablishment varying

4

-

mosaics’ comprising fallow and and fallow comprising mosaics’ HELEN. JONES HELEN. Lullula arborea Lullula ia, Norwich, NR4 7TJ, UK 7TJ, NR4 Norwich, ia, K’s largest K’s largest r

to woodland - cultivated subplots over over subplots cultivated 2 lowland grass lowland

& PAUL M. DOLMAN & PAUL

abundance ite thisarticle as provided byUniversityofEastAngliadigitalrepository

and across all across and

, UK , , -

we heath. heath.

brought toyouby - 1

CORE This article isThis article by protected copyright. All rightsreserved. numbers heath replicated extensively matters closed swards of importance the Although episodic D nesting ( etal. al. the Although & treatments management to inform attention Siriwardena M Keywords grass within lowland within 67 % and 32 % respectively, Conway % respectively, 32 67 %and eclines

anagement anagement Van Bommel 2004 Bommel Van

Accepted2009 Article

2017 ( c involving involving . . ( ) We The -

Mallord ,

may cultivation)

45 .

(hereafter ‘abundance’) (hereafter declines in some areas haveresumed areas insome declines

: While many While ) . grass management, Conservation

across different habitats habitats different across

assessed m of woodland, m woodland, of Woodlark et al. M

British British be

interventions promote promote ost

102 linked et al.

2007 territories territories

- population underwent a partial recovery during the late 20 late the recovery during partial a underwent population )

which which plots heaths characterised by closed swards. closed by characterised heaths

the effects ofground the effects , where where

’s

2007a

multi studies use observed relationships between and speciesand between relationships useobserved studies , to

population ( Holt Lullula arborea Lullula van den Berg den van

, the totalling 248 totalling historically historically for for - bare through annual ground annual through

it is it is ) in Britain in Britain taxa et al. et

cessation cessation and bare and protected under Annex 1 of the EC Birds Directive (EC 1979). 1979). (EC Directive Birds EC ofthe Annex 1 under protected ,

ground

and and

landscape 2014

recovery

are needed to to needed are have been extensively tested beenextensively have

et al. et

habitat use habitat et al. are are create -

)

open open global population is concentrated in Europe Europe in concentrated is population global of dynamic processes dynamic of ha ,

is known, iti is known, but associated with associated -

, within 2009 disturbance on Woodlark as onWoodlark disturbance

2001 , nor whether disturbance disturbance , norwhether - - d scale experiment experiment scale foraging foraging other other heath, lowland heathland, heathland, lowland heath,

early

and the species the and ) , ,

across four across Border where the species use thespecies where

3 support best practice best support lowland lowland - - , disturbance, can increase Woodlark abundance Woodlark increase can disturbance, 850 successional successional s areas areas

not

ha of grass ha of et al. lowland heathland lowland clear whether clear whether semi (

Bowden 1990 Bowden

treatments, differing treatments,

(e.g. rabbit rabbit (e.g.

in in 2017

is - the natural habitats natural mosaics mosaics classified classified

in farmland andwoodland farmland in - ) heath

, experiments , experiments UK’s treatment treatment an integral part of of part an integral s semi

(

grazing, tu grazing,

taller treatments treatments Buckingham largest largest within within ) , . W as Mallord -

or plantation forestry plantation or natural habitat habitat natural Threatened ( Threatened e

vegetation for vegetation th

habitat habitat examined

lowland heathland lowland

century century or habitat type habitat or lowland lowland have received less receivedless have

in in that

rf/ ( et al. SPEC 2; Burfield 2;Burfield SPEC establishment that that

litter removal, litter removal, et al.

composition composition test multiple multiple test

( 2007b

gras open Conway an territory territory Stanbury

2004

(e.g. (e.g. s - - up up ) . ) .

et

.

This article isThis article by protected copyright. All rightsreserved. represent homogenous bothone 2017 subplots cultivation. time since, and of, invaried frequency half complex deep were Treatments controls uncultivated Across Expe 52°43'N 52°51'N carried was out The study Study site sub complex for controlling ‘ method Accepted Article complex plots - overlapping overlapping rimental design design rimental - cultivated the

, 200 ha) , 200 ha),Bridgham , 3,500 METHODS - - ( in 2016 2016 in would - shallow mosaic mosaics mosaics ing mosaic plots mosaic

se

sites similar cost and

- vegetation type vegetation

, be preferred be and s , - 13 and half first half and and

repeated , ( would in in complex or or

2 66

0 deep 0 shallow Eastern England (Fig. England Eastern two deep

four four

were established were replicate ’ ,

compris - suppor ; year

- but 1 from from - - cultivat

m cultivated

in - ha subplots ha subplots cultivated

within complex within osaic

-

- early while Heath ( Heath old time ,

2 t the year 2015 ing

ha ), ed

plot

- first

2016 cultivation subplots varying in age and inage varying subplots homogenous homogenous

plots (3 plots ) and landscape features landscape and - greatest greatest , 52°44'N 0°83'E, 2017 on the onthe 2017 and complexity and ) 20 20 s

- in early 2015 treated annually

time in 2017, that that in 2017, received 2 received

and 2017

shallow S 1; 1; 3 - -

mosaic cultivat for site details, see sitedetails, for deep increase ,

building up - plots remained remained plots cultivated Stanford M Stanford , - Each c Each

ha ofground ha cultivated maintain s

ed , 150 ha) and Brettenham Heath ( Brettenham ha)and , 150 (for . includ (

annually treated annually in abundance ,

and and in the same location, samelocation, in the omplex treatment details treatment

a ed ) . ing , annually rotational rotational ,

ilitary Training Area (STANTA; (STANTA; Training Area ilitary We

again

3 fallowed ( fallowed

disturbance frequency disturbance 3 2 - - Appendix S1) Appendix disturbance treatment each year, year, each treatment disturbance

mosaic comprised comprised mosaic 6 2 ha shallow a priori a

as

cultivating cultivating ,

- and (2) (2) and

2 cultivated cultivated in area in

mosaic of of mosaic ha ‘ homogeno in 2016 2016 in

- predict cultivated homogenous , see recently ,

. complex and

2 ha each year, but but year, 2 haeach

Appendix one ( ed that Fig. Fig. sub diversifying diversifying u ) three three - s

year and and - plots that plots that 1 cultivated plots’ ) 0°83'E, 0°83'E, ) - , . mosaics mosaics

: (1) (1) : while while plots All All

- 36 S 1 old; in old; in 2

; ha 0°76'E,

)

or or .

( 13 40

as This article isThis article by protected copyright. All rightsreserved. subplots complex For from < wind force ( t In each year, and mapping Territory = distribution Wallis controls heath strata and w information, cultivation type( underlying soil ( areas this site disturbance trees complete complex homogenous, monitoring years, designs and increas days between visits: between days Accepted19.26 Article a or scrub ( scrub or , : ,

n (3) intermediate grass (3) intermediate (1) calcareous (1) ed to 3 ha in 2016 and 2016 3 hain to ed vehicle H ,

Plots and Bridgham and Brettenha Bridgham and and

= P (five groups) groups) (five

used by used = 2.65

4 ha 4 16), 16), < (

s Sheail 1979 Sheail 0. - plots plots

4) mosaic of soil types and grass types soil and of were located in were located 0 hree hree

. and positioned positioned

. Ulex 0 ,

During 1; Longitude, 1; Longitude, P

the same the same treat STANTA,restricting of area’ central‘impact in the =

the impact area impact the ithin ithin 40

europaeus s, 0.62; National Soil Resources Institute, Cranfield University Cranfield Institute, Resources Soil National

mean mean grass

- were we minute minute -

) each mosaic subplot use subplot , constraints of ordnance of constraints

and ( and also H > -

individual individual

heath of any age(here ofany heath distributed distributed 24 100 = 1.23, = 1.23, visit,

- grass

heath 4 ha by 2017. ha by 4 H visits recorded recorded 3

) examined consistently ± 11 ± 11 or control) or .

) indicator plant composition plantcomposition indicator )

m away, away, m = P

otential otential we 4 - - P

7

( heath sd , were made to each to made were heath ages, heath n

. and (4) ancient acid grass (4) ancient and

= 0.87 = 19 m

were included Woodlark recorded ) =

similarly similarly between dawn and 11:00 during during still and11:00 dawn between

, 20). H the number of of the number P followed by walk by followed , eath ( eath , for

often often < but including sufficient un sufficient but including ,

To Potential plot locations were mapped weremapped plot locations Potential respectively 0.

unexploded ordna unexploded 0 account for differences in differences account for Fig. Fig.

01 with respect to respect with v , excluding

egetation plots after ;

S n

as separate registrations separate as 1). 1).

= a

were allocated randomly randomly were allocated 102 registrations 4 ha ‘ 4 C calcareous ;

ontrol plots were ontrol location and behaviour location ha n ing ,

)

but close to but close strata strata . = area area

plot between plot between

before 102 through through

- heath (Table heath L nce precluded nce precluded atitude ) centred on the plot ( plot the on centred (four groups) groups) (four ; but due to ; butdue

grass

on on treatment (Table S1). S1). (Table treatment treated grass treated each

each (95% within16m)

- and and

ments heath treatment extent between between extent treatment ), ( 14

also also plot

,

S2

subplot L . dry mornings dry 2

March and March and ongitude aggregated aggregated Observations Observations ) age since last age sincelast ) ) placing ’

. ’ was not was ) to located s to , Treatments ,

edge and centre edge and (2) young grass (2) young initially scanning initially scanning

-

based on based the heath to four vegetation vegetation four ; multiple multiple whether whether

outer outer ground

( in these in (Latitude, (Latitude, Kruskal 26 ,

Using this Using scattered (Beaufort (Beaufort

: ( : were June areas of of areas and 1 - ) -

-

. H

This article isThis article by protected copyright. All rightsreserved. performance model did reduce not werecombined variables with fixed effect annually to subplot per visits) (over three registrations Wright Schaefer (potential refuge, plot homogenous log 2017 last the during and (2) in 2015 homogeneous classified as treated plots S A resident from Conway occasions these on replicates vehicle to restricted eparate eparate nalysis nalysis

Accepted- Article link,

identity identity

plot identity plot identity at least two at least were all were all

et al.

For For F T - considered fixed effects of effects of considered fixed etal.

or occupied occupied or cultivated, cultivated, erritories were subsequently subsequently were erritories a , but not controls , but not nalyses considered nalyses s parsimony,

as a random factor. arandomfactor. as of of . 2007)

v subplots (2009) s cultivation

complex

separate visits separate as a as , ,

were s on 20 20 s on veh one . complex We We

random factor random

two years(2016 two

available) initial i - cles were - year or two or year mosaic both if: (i) if: recognised recognised - method ) to minimise ) to & of 936 of

Vogel 2000) Vogel

categories categories - abundance abundance mosaics inonly, 2017 mosaics entered parameter estimates were were estimates parameter , but excluded males , butexcluded ) . Distance ,

Generalised Linear Mixed Models (GLMM) Models Mixed Linear Generalised reposition vegetation strata vegetation plot

( t ( 2 categories: reatment Akalike's - . year fallow territories territories

- identified across the threevisits the across identified -

visits (3 in 2015, 10 in 2016, 7in 2017 10in 2016, (3 2015, visits in as covariates as 2017), when 2017),

within within disturbance to to disturbance

per per from plot edge to plot from and Thetford Forest (Fig. S1 (Fig. Forest Thetford and ed

Information Criterion corrected for small sample size, size, smallsample for corrected Criterion Information plot

sub (5 (5

during visit during vegetation strata, treatment, and and treatment, strata, vegetation with with , then homogenous orcomplex homogenous , then )

categories , in , in shallow - - treatment year

(4 (4

. complex a GLMM GLMM a we related the maximum number of of number themaximum we related registration

apparently apparently

categories :

Eurasian Eurasian (1) -

initially initially vs deep vs s

: control, cultivation: control, across all three years (2015 years all three across

to the nearest wood nearest the

- (4

with with mosaics had accrued (though only inonly (though hadaccrued mosaics maximise coverage maximise categories s drawn drawn Stone ),

similar - (on, or singing above orsinging (on,

cultivation Poisson and ;

, - potential population source, source, population potential

year curlew in for eachyear for ,

and : to interact to interact

error first

( ,

; affecting affecting 4 ; 2 or 2 ) with land (ii) their (ii)

and vegetation strata, and vegetation Burhinus oedicnemus Burhinus - - - time , method mosaic thereafter mosaic

that 3 sub

.

Po

(> categories -

, i briefly - cultivated, cultivated, 0.5 ha) 0.5

sson error and error sson treatment treatment incorporated incorporated following following combination combination - , 2017

the plot) x

treatment with a with a

;

) with a with ,

with

) ,

ll ;

This article isThis article by protected copyright. All rightsreserved. disturbance distinct combined grass models occupied 2016and2017, In 2015, RESULTS ( resid modelled in confidence the models coefficients 2AICc within lay one model best combinations simplification present categories within possible, where simplified models categories 2;Burnham ≤ increased or AICc, eitherreduced, AcceptedParadis Article package if if - heath Δ

; that that (2015 <

AICc AICc every Table S3) Table et al.

plots plots 2

calcareous with intermediate and ancient and ancient intermediate with calcareous were not not were were Δ ,

( treatment incorporated ‘MuMIn’ ‘MuMIn’ unconditional standard errors standard unconditional AICc retaining (difference in AICc in (difference tervals , -

were examined were 2004 year 2017 and2016 2017 for each for analysis, uals were examined by Moran’s I Moran’s by examined were uals were within 45 m of woodland. ofwoodland. 45m were within examined examined , accounting for theirAkaike weights for , accounting . Subsequent simplification simplification . Subsequent ; ) . thus combined thus

combined combined

of the model the of All models were run in R wererunin models All ( ancient acid ancient Barton 2018 Barton s: 10 cultivation

all within within , , we used multi used , we 25 other other -

2017) using using

in 2015 in and )

the set of of the set relative to all relative 2016 - abundance abundance averaged parameter parameter averaged

) variables as distinct as - categories categories 39 .

package method supported supported C andidate

- - territories were associated with with associated were territories 2017 2017 c , -

andidate models andidate model inference to inference model and 95% confidence intervals intervals confidence and 95% .

x homo ‘lme4’ abundance abundance abundance Next, t Next, ) of 2016 of analys

, whilst simplification in 2017 subplot models models subplot in2017 simplification , whilst ( would would other candidate models candidate other R Core Team 2015 Team R Core Initial s Initial

combining variables , separately for each year, each for , separately &

Anderson 2002 Anderson ( e reatment genous/c Bates acid grass acid - s have have 2017

implification (following Burnham Burnham (following ( 2015 did not not did and and model

were

calcareous with intermediate and young young and intermediate with calcareous et al. be abundance abundance -

2017 2017 comprising 2017 en omplex

and sub and

- deemed to deemed s span heath

estimate

2017 confounded with year confounded as ) . )

sub

and

. complex of vegetation strata vegetation of

First, v First, zero. zero. - )

mosaic ( . - plot

r treatment treatment was > was 2016 M models plots, respectively plots, etaining young young grass etaining

all possible possible all model odels odels across across

Spatial autocorrelation autocorrelation Spatial

be be models using using egetation strata strata egetation & Andserson & - - mosaic

, simplified to treated to simplified , 2017 2. supported combined were accepted as were accepted - W averaged averaged the the those categories ) here .

T and subplot subplot and s variable variable reatment reatment package were not were not .

competing competing

more than more

Following Following in in

2002) if

; all ground all abundance

a

ll - were

heath ‘Ape ,

using using of ’

as -

This article isThis article by protected copyright. All rightsreserved. not supported Table model 2017 strata treatments considered in factornot correlated I= 0.04, Moran’s significant; analyses 0. thisdistance plots double to compared younggrass or intermediate the 2016 from models (2015 in averaged (Table S5). the twoyears this model) in combined were categories treatment S5) & Table (Fig. 2 closer towoodland on higher andwas year, with increased abundance 2017) as ‘recently S3) controlvs. 1

Accepted8 Article models subplot of2017 . Simplification –

. S6 model (where there was no spatial autocorrelation) nospatial was there (where model 0.

(Table S2) (Table (Table S4) (Table 53 In 2017 Multi - there were there ), and on calcareous, intermediate and ancient acid grass acid ancient intermediate and calcareous, on ), and were spatially autocorrelated ( were spatiallyautocorrelated cultivated’ ;

(though (though control

and controls were distributed randomly distributed were controls and

controls - model inference was undertaken for for undertaken was inference model ) - . 2017 model 2017 ,

Model residuals were not autocorrelated not spatially were residuals Model , Woodlark were recorded were Woodlark ,

and and showed showed ,

models models 0.0 ) several candidate models candidate several , and

effect (treatment 9 , 95% CI , 95%

Altho

- from woodland woodland from one more 2017 and2016 2017

retain showed showed s -

P heath, of treatment and distance to woodland w towoodland distance and treatment of

- = 0.009 ugh vegetation strata and distance to Thetford Forest were included included were Forest Thetford to distance and strata vegetation ugh year

registrations registrations 0. , ing

01 0. - the 18 a higher abundance on on abundance higher a old 59

– complex . 0. )

, , Multi m from m

modelling

with al 95% CI 95% 17 suggesting some variation attributable to aspatially to attributable some variation suggesting - though only in 2017, where Moran’s I was small but smallbut Iwas Moran’s in where only 2017, though

(36 m) 2017), neither 2017), on ; combined Fig. 3).Fig. - two

model inference from from inference model - in 21 mosaics/

where where woodland 0.

recently / -

40 37 h ; year

nevertheless ad M both all treat all –

, complex in the landscape thelandscape in

0. but odel averaged residuals odel averaged

first < a lower abundance abundance a lower - old fallows (hereafter‘fallows’ fallows old 81 homogenous homogenous 2 - abundance with no support for a difference be difference a for nosupport with cultivated

effect was supported effect was ; - Δ . (median

time

control AICc AICc ed treatment treatment - mosaic plotsmosaic

plots -

, we consider consider , we (Table (Table S4). with annually with , -

. plot 0.1

heath heath

than 2016

( analyses compared to controls compared were similar and 5 plots - ere , woodland woodland

95% CI 95% - fallow subplots fallow

( 2017 ( balanced across vegetation vegetation across balanced . cultivation treatment

F consistent with consistent T or 2015 or in 2017, 2017, in he

from the from inference inference (2015 - cultivated

was similar ( similar was (Table S5). P S5). (Table best supported 0.0 ; distance) ∆ 3 - - AIC 2017 and2016 2017 – 2017 , on calcareous, calcareous, on -

; 0.2 method 0. Table S3). 2015 c robust, as robust, 36

= 1. (Fig. 4 & (Fig. 7 (hereafter (hereafter , ), whilst ), whilst , 95%

) the

, redictions though though and plots plots and

- tween 9 2017 ;

was

Table Table 2016

CI

- -

This article isThis article by protected copyright. All rightsreserved. birds andshowed Woodlark, for ecotones forest to woodland. plotsclose for vegetation woody was territories natal dispersal discover ( which showed Switzerland, preference for to recent recent other and complex 2017) numbers 1996 shown research has woodland, respond Woodlark landscape replicated anextensively Through D Arlettaz Arlettaz Accepted Article ISCUSSION

similar , . , Mallord

- This abundance was greater on greater was abundance cul C T toour Contrary ed - et al. he onsistent with with onsistent have have cultivation in a in a cultivation

tivation

was not was

and and might be because be might treated plots plots treated between treatments between increase increase - mosaic mosaic

et al. 2012) been influenced experimentally influenced been the barer ( select e.g. up to 11 km; Bowden Bowden 11 km; to up e.g.

in a matrix amatrix f of in

benefit ed due to due

2007b Woodlark Woodlark . plots from from ed positively to positively

a priori a evidence from Iberia from evidence recently recently ( matrix ofgrass matrix

visualised by Fig. Fig. by visualised ted ’ the increasing siz the increasing , , 20

Arlettaz c compared to compared

Schaefer and Vogel (2000) andVogel Schaefer . 50% ground 50%ground . :

15

Woodlark Woodlark (i) require - prediction, prediction, . 2017 - both cultivation both a

- cultivated subplots cultivated

cultivated subplots cultivated

llows (complex llows both all

et al. et

ground

was was

bare ‘ - shallow and deep and shallow (

heath (homogenous plo (homogenous heath Sirami

controls 2012 & attributed to attributed vegetation vegetation S3 - w e and complexity of the complex ofthe complexity e and

open Green 1992 - - , hen scale experiment scale disturbance )

where where ,

) through mechanical interventions mechanical through

consistent with adult fidelity and the known scale of scale known the fidelity and adult with consistent , but as far as we are aware we faras are as but , et al.

-

fly towards forest when disturbed, disturbed, when forest towards fly -

all methods methods , areas areas mosaics) offers little by way of additional resource resource additional of littleoffers byway mosaics)

but but

treatment combinations combinations treatment

within complex colonisation of colonisation 2007 ( cover is optimal cover is Fig. S2) the

explored the ecological function offield function the ecological explored for foraging for cumulative cumulative ) .

- treatments We are confident confident are We se ) cultivated cultivated create , our

treatments treatments

is consistent with is consistent ,

we we

results demonstrate results ts) d

have have suitable .

colonisation colonisation

( - previously open habitats by by habitats open previously complex Within Bowden 1990 Bowden treatments , mosaic treatments mosaic

prefer for foraging foraging Woodlark for demonstrated that demonstrated did ,

foraging habitat, and and habitat, foraging

this - this is the first time isthefirst this not differ not red mosaic were available

a study from a study ’ accumulation of accumulation plots closer to plots as

. and

,

individuals Sitters s, as s,

stating - d mosaics, mosaics, ‘

from each each from homogenous a preference a preference .

abundance abundance Previous Previous

et al. et

on on (2016

their -

(ii) -

This article isThis article by protected copyright. All rightsreserved. AcceptedChamberlain) assistance DIO Organis Infrastructure Defence from with support RSPB and ground scarce species this management deter Articledeep cultivated or intervention) necess g and disturbance, prescription that ground suggests Our experiment recommendations Conservation open of woodlark anti of closer examination , Sheep Enterprise Sheep other potential beneficiaries beneficiaries potential other ary - - disturbance Although heath and clear and heath -

Natural England England Natural territories . Cranfield U . Cranfield ( al

for Woodlark for though methods which retain bare ground for longer may require less less may require forlonger ground bare retain which methods though , regardless of regardless of ,

and one anonymous reviewer reviewer anonymous and one

( Dolman

Woodlark Woodlark are unclear are - iven iven cultivated)

, prescription

but the habitat of that species of habitat but the , NE , niversity the

et al. et - fell fores fell

, - within other lowland grass lowland within other (NE) (NE) ir predation strategies it became clear that that clear it became strategies predation Richard establishment method establishment

responded positively responded preference preference

, 2012 . al Interventions

funded this work this funded provided soil data underlicence data soil provided though though

until the until the try; however, try; however, ,

that prefer open habitats habitats preferopen that Evans, Evans, Pedley - disturbance could could disturbance autoecolog for recently for wider results of our experiment areavai experiment ofour results wider STANTA bird group bird STANTA

et al.

for helpful comments on on comments helpful for should be be should th

2013 through the the through

to all eir association with woodland edge with woodland association eir ation

(complex ical . - ’ cultivated subplots cultivated

- In ) heaths . W

treatments information information within within

(DIO Britain, Britain, represent e

thus ) - . ( mosaic or homogenous; shallow orhomogenous; mosaic , and Breaking New Ground BreakingNew and Action for Birds inEngland forBirds Action e.g. Stone e.g.

Since all Since c Dominic Ash and Ian Levett for Levettfor Ian Ash and Dominic . Woodlark are regarded as a species asa areregarded Woodlark .

45 advise caution in in caution advise

We are grateful to We aregrateful ,

an important important an m woodland of the indicates forest edge is not a factor of afactor is not edge forest an earlier version an earlier treatments treatments multi , - curlew, Johnston 2009 Johnston curlew, annual

- it it will taxa consequences of of consequences taxa - cultivation cultivation conservation conservation using lab involved annual involved , but , but

benefit

frequent frequent the

le. is

a single a single this

editor . We . of this paper this of important

p rogramme

many may be may may

thank thank -

(Dan ) .

.

, .

This article isThis article by protected copyright. All rightsreserved. M. P. Dolman, R.J.& Fuller, Holt,C. A., H.L. J.&Mossman, C. M., Panter, P. Dolman, J. Spencer, A.& Drewitt, Eaton, M., I., Henderson, Wotton, G., S., Conway, D. R. Anderson, P. & K. Burnham, F. Van Bommel, I.& Burfield, A. J. Rook, P. & D. L., Atkinson, Buckingham, R.E. R.&Green, C.G. Bowden, C. Bowden, I. R. Hartley, J.W.& G.,Redhead, I. Henderson, A., Border, J. S. Walker, & B. M., Bolker, D.,Maechler, Bates, Barton, K. Mosimann L., M. Maurer, R., Arlettaz, R Accepted Article EFERENCES woodland regeneration woodland inconservation. mismatch a identifies and regional priorities breeding information status, research. current A reviewof report.Sandy. RSPB Unpublished forests. andSandlings the Thetford in plantations plantations. Whinchats at:https://cran.r available 2018). September 23 (accessed Woodlarks. grou patchy create practices cultivation New2012. vineyard

Multi

1990. Selection of foraging habitats by Woodlarks ( Woodlarks by habitats offoraging Selection 1990.

Cambridge: Birdlife International. Birdlife Cambridge: - Model Inference. available at: https://cran.r at: available Model Inference. Saxicola r Saxicola - J. Ornithol. J. Appl. Ecol. Appl. J. theoret ic approach, ic approach, ubetra Lullula arborea Lullula

2004. 2004.

153:

in winter and summer. summer. winterand in 1992. The Ecology and managemen and TheEcology 1992.

27:

- 2002. 2002. project.org/web/packages/lme4/ (accessed 3 May 2018). 2018). 3May (accessed project.org/web/packages/lme4/

at territory and landscape scales. and landscape territory at Birds in Europe: population, estimates, trends and conservation conservation and trends estimates, population, Europe: in Birds 229

410. Ibis - Kampe, P., Nusslé, S., Abadi, F., Braunisch, V. & Schaub, & M. V. Schaub, Braunisch, F., Nusslé,S., Abadi, P., Kampe,

- 2014. Exclusion of deer affects responses of birds to ofbirds affects responses ofdeer Exclusion 2014. Model selection andmulti selection Model New York: Springer. York: New 238.

146:

in Britain in 2006. Britain in in

2004. Testing solutions ingrass solutions Testing 2004.

163 20

Linear Mixed Linear

12. The biodiversity audit approach challenges challenges audit approach biodiversity 12. The - 170. Ibis

156:

- project.org/web/packages/MuMIn/ project.org/web/packages/MuMIn/

-

2017. Habitat selection by breeding breeding by Habitatselection 2017. Effects Models using 'Eigen' and S4. S4. 'Eigen' and using Models Effects Bird Study Bird Lullula arborea Lullula

116 - model inference: a practical apractical inference: model t of Woodlarks on pine onpine t Woodlarks of nd vegetation, favouring favouring vegetation, nd - 131. J. Appl. Ecol. J. Appl. Ibis

56: 159:

2009. The status of of status The 2009.

- 310 dominated landscapes: landscapes: dominated

) nesting in pine in ) nesting 139 -

325. 49: - 151.

986

- 997.

This article isThis article by protected copyright. All rightsreserved. Accepted G.R. & Bowden, C. N. T., Cowie, M. R.Wright, Hoblyn, A., R. P., Fuller, J., H. Sitters, Dodd, N.A.& S. A., Calbrade, J. A., Vickery, Q. G. D.K., Anderson, M., Stevens, G. Siriwardena, &Martin, J. L. Brotons, Sirami, C., Sheail, J. B. & Vogel, T. Schaefer, Team. R Core ArticlePedle K. & Strimmer, Paradis, J. Claude, E., J. W. Sutherland, F.& A. M., Brown, P. Dolman, J.W., Mallord, J. W. & Sutherland, A. M., Brown, P. Dolman, J.W., Mallord, Johnston, A. y, S. M., Franco, A. M. A., Pankhurst, T. & Dolman, P. M. P. Dolman, T.& A.,Pankhurst, A.M. Franco, y, S.M., Bird StudyBird Woodlark two large from birds: evidence farmland of populations breeding on winterseedfood ofsupplementary Theeffect 2007. plot. census to From landscape the grass factors. possible Computing. forStatistical R Foundation 182. experiment. taxa A multiple biota: forheathland networks ecological language. size inaground population to disturbance and productivity? fornestsurvival consequences Woodlarks ofCambridge. University thesis. Cambridge: PhD populations.

1979. Documentary evidence of th evidenceof Documentary 1979.

2009. Demographic analsis of the impact of conservation action on Stone on action conservation impact of the of analsis Demographic 2009.

2015. 2015. - heaths of Breckland. Breckland. heaths of

Bioinformatics Lullula Lullula arborea 43: Lullula arborea R: a language and environment for statistical for environment and language a R:

172

2000. Why do Woodlarks need field Woodlarks Whydo 2000. J. Ornithol. - 187.

20:

in Britain: population trends, distribution and habitat occupancy. occupancy. habitat and distribution trends, population Britain: in

2007. Vegetation and response to land abandonment: abandonment: toland response songbird and Vegetation 2007. breeding on heathlands in southern England: are there are England: southern in heathlands breeding on

141:

289

2004. APE: Analyses of Phylogenetics and Evolution in R Evolution in and ofPhylogenetics APE:Analyses 2004. - J. Biogeogr. scale experiments. scale experiments. Divers. Distrib. Divers.

- 335 290. e changes in the use, management and appreciation of andappreciation management use, inthe e changes - 344.

-

nesting . passerine. nesting 6:

277

13 : - Bird Study Bird

292. 42 J. Appl. Ec J. - forest ecotones? ecotones? forest -

52. 2007a. Nest 2007a.

2013. Physical disturbance enhances enhances disturbance Physical 2013.

2007b. Linking recreational recreational Linking 2007b.

computing. Version 3.2.2, 3.2.2, Version computing.

54: ol.

J. Appl. Ecol. Appl. J.

44: 307 - site characteristics of characteristics site

920 - 314. Biol. Conserv. Biol. — - 932.

An analysis of of An analysis

44:

1996. The 1996.

185 - curlew curlew - 195.

160: Vienna:

173 - This article isThis article by protected copyright. All rightsreserved. Table S6 AcceptedTable S5: Table S4: Table S3: Table S2: Table S1: Figure Figure Figure S1: S2 Appendix S1 Appendix ofthisarticle: version inthe online be found may Information Supporting Additional Article INFORMATION SUPPORTING M. P. Dolman, J.& W. R.Sutherland, Hoblyn, A., J., Wright, L. J &R. H. W. Rose, Langston, T., R. R. J.M., Clarke, Bullock, J.L., L. Berg, van den A. Brown, A., Stanbury, S3 S2 323. Woodlarks size. population and fragmentation type, habitat ( warbler Dartford bythe selection R. Gregory, :

: : S Abundance models Candidate S Plant Model simplification

S T Complex ubplot model ubplot trata trata tudy site tudy erritories perplot erritories

: : T Site

reatments indicators

information - Lullula arborea Lullula arborea

mosaic subplots mosaic 2017. The risk of extinction for birds in Great Britain. GreatBritain. birdsin for extinction Theriskof 2017.

models , Eaton, M., Aebischer, N., Gillings, S., Hearn, R., Noble, D., Stroud, D.& D.,Stroud, R.,Noble, Hearn, S., N.,Gillings, Aebischer, M., , Eaton,

-

year

in traditional and recently colonized habitats. habitats. colonized and recently in traditional

Sylvia undata Sylvia ) in Dorset, England: the role of vegetation vegetation roleof England: the Dorset, ) in

Biol. Conserv. Biol.

2007. Reproductive success of of success Reproductive 2007.

101: Br. Birds Br.

217 - 228. . Bird study Bird

2001. Territory Territory 2001. 110:

502 -

54: 517.

315

- This article isThis article by protected copyright. All rightsreserved. Accepted estimates SeeFig respectively. years, consecutive one 0:first plot/subplot; each of frequency age/disturbance the denote Numbers Figure Article - year 1 .

- Development of homogenous and complex homogenous of Development old fallow; 2:two fallow; old

for

each each complex - - year mosaic mosaic - old fallow; x2 and x3: annually x3: x2 and fallow; old sub ure - treatment in2017. treatment

S2 for example for - mosaic plots over three years (2015 threeyears over plots mosaic

photographs

- cultivated

and % bare ground ground % bare and

in each of two and three three two and each of in - time - cultivated; 1: cultivated; - 2017).

This article isThis article by protected copyright. All rightsreserved. Accepted and 2017 2016 of2015, ineach controls treatments and ArticleFigure 2 .

Mean Mean (± se ) number

of Woodlark territories per plot ( perplot territories Woodlark of

.

n

= 102) forground = 102) - disturbance disturbance

This article isThis article by protected copyright. All rightsreserved. Accepted respectively. intervals, confidence and95% means predicted multi on based are Predictions heath. nearest wood treated(darkgrey across Figure 3 Article .

Predicted Woodlark Woodlark Predicted land ,

f or (a) calcareous, (a)calcareous, or , n = 66 Lullula arborea Lullula ) and control (lightgrey control ) and - intermediate and young grass young and intermediate model inference (Table S5) (Table model inference

abundance abundance , n = in 2017 (the final year of the experiment) experiment) ofthe year (the final 2017 in 36 Circles

) plots in relation to distance to the to distance into relation plots )

.

L ines and shading represent represent shading and ines -

show individual data points. data individual show heath and (b) ancient grass acid (b) ancient heath and

-

This article isThis article by protected copyright. All rightsreserved. Accepted represent Bars ± from subplots Figure Article 4 .

Mean number of Woodlark registrations per sub per registrations ofWoodlark number Mean 21 occupied complex 21 occupied se , unfilled circles show individual data individual circles show , unfilled - mosaic plots in 2017 2017 plots in mosaic points (11 deep - treatment (filled symbol) across 84 across (filled symbol) treatment

.

- cultivated, 10 s cultivated,

h allow - cultivated) .