NunezRegueiro Et Al2015-BioCon (1)

8/16/2019 NunezRegueiro Et Al2015-BioCon (1)

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Biological Conservation 187 (2015) 19–2

Contents lists available atScienceDirec

Biological Conservation

journal hoe!age"###$elsevier$co%locate%bioco

Spatial patterns of mammal occurrence in forest strips surrounded by

agricultural crops of the Chaco region, Argentina

Mauricio M. Núñez-egueiro a,&, !yn "ranch a, obert #. $letcher #r. a, %usta&o A. Mar's b,

(nri)ue *erlindati b, Andr+s 'lamo c

a ni&ersity of $lorida, *epartment of ildlife (cology and Conser&ation, //0 Ne1ins-2iegler 3all, %aines&ille, $! 456//-0740, nited States b ni&ersidad Nacional de Salta,

$acultad de Ciencias Naturales, A&. "oli&ia 8/80, Salta 7700, Argentina c ni&ersidad Nacional de Salta, $acultad de Ciencias Naturales, Conse9o Nacional de :n&estigaciones

Cient;ficas y +cnicas, A&. "oli&ia 8/80, Salta 7700, Argentina

a r t i c l e i n f o

Article history<

ecei&ed /= No&ember 50/7

ecei&ed in re&ised form /> March 50/8

Accepted / April 50/8

?ey1ords<

$orest strips

$ragmentation

Species ecological traits

*eforestation

Chaco forest

"ayesian occupancy models

a b s t r a c t

*eforestation is a ma9or cause of biodi&ersity loss, and the predominant factor dri&ing deforestation is e@pansion

of agriculture. A ey step to1ard successful conser&ation in agricultural areas is ma@imizing biodi&ersity &alue of

remaining forest. :n subtropical and tropical regions, forest often is left in narro1 strips bet1een agricultural

fields under the assumption that biodi&ersity is sustained. e e@amined use of forest strips and continuous forest

by medium and large-sized mammals in Argentine Chaco 1ith camera trapping and hierarchical "ayesian zero-

inflated occupancy models and assessed ho1 use related to ecological traits of species. Almost =0B of the species

cited for our study area 1ere not detected or 1ere detected in less than /0B of the sampling units. en of the 54

species that occurred in the area 1ere absent from strips or 1ere detected most fre)uently in continuous forest,

including all large-bodied species and forest interior specialists. !o1 occurrence of mammals in strips and in

continuous forest raises ma9or concerns related to long-term persistence of mammals in Chaco. nder current

de&elopment policies, agriculture 1ill continue to e@pand in this region, further threatening the second largest

forest in South America. Alternati&e configurations for the forest-agriculture landscape, as 1ell as synergism

bet1een landscape configuration and other threats, need to be e&aluated and incorporated into policy if the rich

mammalian fauna of this region is to be conser&ed.

ublished by (lse&ier !td.

/. :ntroduction

"ecause of massi&e con&ersion of forested habitats to agriculture,

many pre&iously forested landscapes comprise large e@panses of crops

interspersed 1ith forest patches and strips D3a1es et al., 500EF.

nderstanding and predicting ho1 1ildlife responds to different

configurations of forest in agricultural landscapes is essential for

de&elopment of effecti&e conser&ation policy. *eforestation is a ma9or

cause of loss of biological di&ersity, and the largest factor dri&ingdeforestation is e@pansion of agriculture to supply an increasing

demand for food, biofuels, and other agricultural products D$AG,

50/5F. $orest buffers are often left along ri&ers and streams to reduce

agricultural run-off and protect 1ater )uality D Naiman and *escamps,

/>>=F, and upland forest strips are fre)uently retained around

agricultural fields as 1indbreas and to pre&ent soil erosion and spread

of pests and fire D(risson et al.,

500/F. Although upland forest strips liely are not as 1idespread as

riparian buffers, these forest strips occur in agricultural landscapes in

tropical and subtropical regions across the planet He.g., Argentina

DSeghezzo et al., 50//F, "oli&ia D!. "ranch, pers. obs.F, :ndia DSreear

et al., 50/4F, and araguay D(risson et al., 500/FI. $orest strips and

their configuration De.g., 1idthF historically ha&e been dictated by

go&ernment policy to benefit agriculture and, more recently, 1ith the

additional assumption that these strips maintain biodi&ersity D(rissonet al., 500/J Seghezzo et al., 50//F. his pattern of land con&ersion

thus represents a deliberate landscape design, albeit often a design

1ith little consideration of conser&ation outcomes D$ig. /F.

*espite the 1idespread distribution of forest strips in production

landscapes, the assumption that maintaining forest strips around crops

allo1s forest biodi&ersity to persist in agricultural landscapes remains

largely untested D!aurance and !aurance,

& Corresponding author at< ni&ersity of $lorida, School of Natural esources

and (n&ironment, //0 Ne1ins-2iegler 3all, %aines&ille, $! 456//-0740, nited

States.

(-mail address< mnuregKyahoo.com DM.M. Núñez-egueiroF.

h ttp


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M.M. Núñez-egueiro et al. L "iological Conser&ation /E= D50/8F />56 50

/>>>J "audry et al., 5000F. $orest strips ha&e been e&aluated in a

limited range of ecological systems, principally temperate systems and

to a lesser degree in 1et tropical forests, and most studies ha&e

focused on small &ertebrates, particularly birds D3a1es et al., 500EJ

ehling and *iemann, 500>F. Conclusions to date are mi@ed. $or

e@ample, studies in Amazonia ha&e found greater acti&ity of large

terrestrial mammals in strips than in ad9acent forest D"arlo1 et al.,

50/0F, similar abundances of frogs and small mammals in remnant

forest strips and ad9acent continuous forest D de !ima and %ascon,

/>>>F, and depauperate communities of birds and primates in strips as

compared to continuous forest D!ees and eres, 500EF. o inform

policy and design agricultural landscapes that incorporate

conser&ation goals, understanding of conser&ation &alue of strips

needs to be e@panded to a greater range of geographic

regionsLecological systems and a broader set of ta@a.

esearch on forest strips, and forest fragmentation in general,

suggests that species responses to the forest strips may be di&erse and

possibly idiosyncratic D3enle et al., 5007F. nderstanding of ho1

species traits De.g., life history attributes and dietary and habitat

re)uirementsF influence the ability of species to use forest strips may

impro&e prediction of impacts of forest con&ersion to croplands

surrounded by forest strips and facilitate design of landscapes that

promote species persistence. Such trait-based approaches ha&e

pro&ided insights into &ulnerability of species to climate change,

habitat fragmentation, and hunting D*e&ictor et al., 500EJ *iamond et

al., 50//J hornton et al., 50//J Chessman, 50/4F. eproducti&e rate,

determined by traits such as age at first reproduction and litter size,

influences ability of species to persist in the face of human-induced

mortality factors such as hunting, as 1ell as population reco&ery

follo1ing a decline DAltrichter, 5008F. "ody size often correlates

negati&ely 1ith reproducti&e rate, but also can contribute to

&ulnerability because species 1ith large body size re)uire large

)uantities of food and space that may be not a&ailable in human-dominated landscapes Dhornton and $letcher, 50/7F. Also, these

species often are preferred as game Deres, 500/F. 3unting is a

significant threat throughout tropics and subtropics, and e@posure of

species to hunting often increases 1here forests are fragmented Deres,

500/F. ulnerability of species to con&ersion of continuous forest into

remnant forest strips also may be in&ersely related to ability of species

to e@ploit multiple habitats and food resources. A general pattern

emerging from many studies is that specialist species are more liely

to respond ad&ersely to a &ariety of global changes than are generalist

species D*e&ictor et al., 500EF. eduction in abundance or total loss of

specialist species can result in a shift to communities dominated by

generalist species, resulting in a decrease in functional di&ersity and

biotic impo&erishment in the form of functional homogenization of the

community DGlden, 5006F.

$ig. /. DaF ro&ince of Salta, Argentina, and satellite image of modified Chaco forest including remnant forest strips Ddar stripsF imbedded in 47,000 ha of agricultural land Dlight s)uares, each 500

haF, and ad9acent continuous Chaco forest in Salta, Argentina. Satellite image do1nloaded from %oogle (arth. DbF Aerial photograph of forest strips in a recently deforested area in the Chaco region.

Aerial photograph courtesy of %reenpeace Argentina.


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M.M. Núñez-egueiro et al. L "iological Conser&ation /E= D50/8F />56 5/

e e@amine the conser&ation &alue of forest strips for medium

and large mammals in Argentine Chaco 1ith the goal of informing

on-going land use planning that 1ill largely determine the future of

forests in this region DSeghezzo et al., 50//J i)uer-odriguez et

al., 50/8F. ropical dry forests and sa&annas, such as Chaco, are a

high conser&ation priority 1orld1ide, because these regions ha&e

suffered e@tensi&e habitat con&ersion D3oestra et al., 5008F. he

Chaco forest of Argentina, "oli&ia, and araguay is the second

largest forest in South America after the Amazon forest, and 60

percent of this forest occurs in Argentina Di)uer-odrigez et al.,50/8F. *i&ersity of medium and large mammals in Chaco ri&als

that of tropical forests in South America, and endemism is high

DMares, />>5J edford et al., />>0F. $orest co&er in Chaco

remained relati&ely intact until recently 1ith e@tensi&e land uses

focused on cattle ranching, charcoal e@traction, and selecti&e

logging D%asparri and %rau, 500>F. :n the last t1o decades,

e@pansion of modern agribusiness has rapidly accelerated forest

con&ersion in this region D3ansen et al., 50/4F. :n northern

Argentina current en&ironmental norms re)uire that forest strips at

least /00 m 1ide be left surrounding agricultural plots 1hen forest

is cut D54 4= ha of forest strips for e&ery /00 ha of deforested

landF, resulting in huge e@panses of commercial crops, particularly

soybeans D%lycine ma@F, di&ided by a grid of forest strips D $ig. /,

%inzburg et al., 50/5F. An increasing number of studies document patterns of land-co&er change in Argentine Chaco, but field studies

are scarce and &ery little is no1n about impacts of this process on

biodi&ersity or, more specifically, the &alue of these modified

landscapes for 1ildlife Di)uer-odriguez et al., 50/8J eriago et

al., 50/8F.

e e&aluated use of forest strips in Argentine Chaco by

medium and large mammals Dbody 1eight O / gF and assessed

ho1 this use relates to their ecological traits. e e@pected

composition of mammals to differ in forest strips and continuous

forest, and that the most &ulnerable species Di.e., species that are

more liely to be absent from forest stripsF 1ould be those 1ith

lo1 reproducti&e rate, large body mass, narro1er habitat and

dietary breadth, and species that are se&erely hunted. his 1or not

only supports a broader understanding of effects of habitat loss andcreation of forest strips, but also documents the critical situation for

1ildlife in Chaco, 1hich has the 1orldPs highest deforestation rate

D3ansen et al., 50/4F.

5. Materials and methods

5./. Study area

Gur study 1as conducted in Chaco forest of Salta ro&ince,

Argentina. he study area D>000 m5J center, 5705>04/00S,

6408505400F is dominated by soybeans and pasture 1ith strips of

forest 80/00 m 1ide bet1een all agricultural plots, surrounded by

large blocs DO/000 m5F of continuous forest D$ig. /aF. ithin the

forest, traditional land-use practices of cattle ranching, charcoale@traction, and selecti&e logging continue around small homesteads

Done or t1o familiesF scattered throughout the forest D Altrichter,

5008F. Subsistence hunting is common, occurring opportunistically

during other acti&ities and during targeted hunts into the forest

e&ery one-t1o 1ees. eople from neighboring to1ns also

occasionally hunt in the area.

Nati&e &egetation in our study area is representati&e of dry

Chaco forests, dominated by )uebrachos HSchinopsisi lorentzeii

DAnacardiaceaeF and Aspidosperma )uebrachoblanco

DApocynaceaeFI and accompanied by "ulnesia sarmientoi

D2ygophyllaceaeF, rosopis alba, rosopis nigra, Anadenanthera

macrocarpa D$abaceaeF, 2iziphus mistol DhamnaceaeF,

hyllostyllum rhamnoides DlmaceaeF, and Callicophyllum

multiflorum DubiaceaeF DMorello and Adamoli, />=7F. "ased on

published accounts of terrestrial mammals, 40 species O / g

should occur in our study area DMares et al., />E>J Cane&ari and

accaro, 500=J allace et al., 50/0F.

5.5. Sampling design

$rom May 50/0 to #anuary 50//, 1e sampled occurrence of

mammals in /5 transects placed in strips of remnant forest bet1een

agricultural fields and e@tending into ad9acent continuous forest.Sampling occurred during dry season because roads are largely

impassable during 1et season. sing %oogle (arth, 1e identified

all strips of forest in our study area that 1ere at least /6-m long.

Most of these strips 1ere created bet1een />>8 and 5007. e only

considered strips at least E-m apart to assure biological and

statistical independence. his distance is larger than home range

diameters of all but the largest carni&ores Dpuma, 9aguarJ allace et

al., 50/0F. e only sampled strips not se&erely degraded by fire

Di.e., Q/8B of strip area affected by fireF. All strips 1ere /00 m

1ide. o a&oid biases from nearby roads, transects 1ere no closer

than 4 m to any ma9or road. Si@teen forest strips met the

re)uirements described abo&e, and /5 strips 1ere selected

randomly for sampling. Species occurrence data 1ere collected

1ith cameras along transects that incorporated E m of the foreststrip and E m of ad9acent contiguous forest. (ach /6-m transect

1as treated as a bloc in occupancy models, and each camera

1ithin a transect 1as considered a sampling unit Dsee belo1F.

$i&e cameras D"ushnell rophyCam, "ushnell Corporation,

G&erland ar, ?S, SAF 1ere placed in each strip of forest, and 8

cameras 1ere placed in continuous forest along each transect. :n

continuous forest, the first camera 1as placed at the limit 1ith the

forest strip and then cameras 1ere placed at inter&als of /.6 m. :n

forest strips, a camera 1as placed e&ery /.6 m beginning /.6 m

from the edge of continuous forest. Cameras 1ere placed in the

nearest appropriate location to designated points Dal1ays Q 8 m

a1ayF, including game trails, den sites, and other areas containing

animal sign. e placed the camera sensor appro@imately /050 cm

off the ground so that smaller species could not a&oid detection by1aling under the sensor. Cameras remained at each sampling

point for /6 days follo1ing hornton et al. D50//F, 1ho studied

similar species. !ogistics of sampling o&er our entire study area

during the dry season precluded a longer sample period. All

cameras on a transect 1ere sampled simultaneously. e recorded

presenceLabsence for each species o&er e&ery 7-day inter&al to

create a series of repeat detectionL non-detection data for modeling

detection probabilities DMac?enzie et al., 5005F. %enerally, 6=

days occurred bet1een deployments at different transects.

5.4. Species traits for &ulnerability analyses

$or the 40 mammal species cited for our study area, 1e determined

trophic le&el, body mass, dietary and habitat breadth, litter size, and

age at first reproduction from field guides and recently published

studies. ulnerability to hunting 1as assessed by inter&ie1ing 5=

residents from homesteads in forest surrounding our transects.

:nformants grouped species as< / R rarelyLne&er hunted or illedJ 5 R

occasionally hunted or illed, but not a preferred game species or

acti&ely persecuted speciesJ 4 R often hunted or illed. *etailed

procedures for deri&ing species traits and correlations among traits are

in Appendi@ A/. "ody size and age at first reproduction 1ere the only

highly correlated traits Dr O 0.60F. Conse)uently, age at first

reproduction 1as omitted from models. 5.7. Assessment of

en&ironmental co&ariates

nderstory density and total canopy co&er Di.e., shrubs and treesF

1ere included in models as potential factors influencing probability of

detection and occupancy. Camera sensiti&ity can be affected by solar


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S!ecies naea S!ecies traitsb 'uber o sites

#ith etections

55 M.M. Núñez-egueiro et al. L "iological Conser&ation /E= D50/8F />56

radiation, 1hich &aries 1ith total canopy co&er Dsee "ushnell

rophyCam 500> manualF. e estimated understory density and

canopy co&er from standardized digital photographs centered at each

camera site D3alcha et al., 50//F. "oth &ariables 1ere highly &ariable

and did not differ significantly bet1een continuous forest and stripsDAppendi@ A5F. An inde@ of local hunting pressure also 1as included

in initial models of occupancy. o obtain this inde@, 1e calculated

density of homesteads around each camera 1ithin an E-m radius

using Arc%:S/0. his distance is the a&erage ma@imum distance

tra&eled per day on foot by hunters in Chaco forest in Salta D !eae,

500>F, and falls 1ithin the range of other &alues reported for Chaco

DAltrichter, 5008J 3ill et al., />>=F.

5.8. 3ierarchical "ayesian occupancy models

Species occupancy 1as analyzed 1ith hierarchical occupancy

mi@ed models 1ith zero inflation accounting for o&erdispersion using

"ayesian inference and history of detections collected from camera

traps Doyle and *orazio, 500EF. hese models incorporate detection probabilities to o&ercome sampling biases related to differences in

species detection that can result in biased estimation of the relationship

bet1een species occurrence and habitat co&ariates Doyle and *orazio,

500EF. 3ierarchical "ayesian models also can incorporate random

bloc effects and, thus, 1ere appropriate for our nested study design

De.g., cameras located 1ithin transects, oyle and *orazio, 500EJ ota

et al., 50//F.

e modeled occupancy for all species 1ith a na&e estimate of

occupancy O/0B of sampling units DN R > species, able /F. hese

occupancy models had t1o components< probability of detection DpF

and probability of occupancy D1iF. :n final models presented here, 1e

modeled p as a linear function of t1o en&ironmental co&ariates,

canopy co&er and understory density. e modeled 1i as a function of

distance along the transect 1here the species 1as detected Dfi@ed

effect, distance hereafterF and transect Drandom effectF because

sampling units 1ithin transects liely are not biologically independent.

he magnitude of the parameter estimate for distance signals strength

of association bet1een distance categories along a transect and

occupancy. Negati&e &alues indicate that a species 1as less liely to

occupy a gi&en site as distance

first reproductionJ see Appendi@ A/ for full description of species traits.

from forest interior increased, and positi&e &alues indicate the opposite

pattern. he full model for each species included additi&e effects of

both en&ironmental co&ariates on p, as 1ell as distance and transect

effects on 1i. e also initially considered other co&ariates De.g., inde@

of huntingF that 1ere discarded because credible inter&als associated

1ith these models o&erlapped 0 Dsee Appendi@ A4F.

*etection and occupancy components of the hierarchical model

1ere analyzed simultaneously using program in"%S called

remotely from program 1ith 5inbugs D%ils et al., />>7J Sturtz

et al., 5008F. $or details on modeling and code, see Appendi@ A4.

5.6. elationship bet1een &ulnerability and species traits

o understand lins bet1een species traits of mammals and their

presence in forest and strips, 1e used t1o approaches. $irst, 1e

di&ided species into three categories. he first category DN*F

comprised all species not detected in our study area that should occur

in the area based on the no1n distribution of these species D Mares et

al., />E>J Cane&ari and accaro, 500=J allace et al., 50/0F. !ocal

forest residents confirmed that these species ha&e occurred in the area

in the near past. he second category D$F corresponded to species that

1ere detected only in continuous forest. !ast category D$ T SF included

species that 1ere either detected in strips of forest and in continuous

forest or only in forest strips Done speciesF. e used classification tree

able /

Number of sites 1ith detections of mammals in continuous forest and forest strips, and life history and ecological traits of these mammals in Chaco forest. he ma@imum number of sites 1here a

species could be detected 1as 8 sampling units in forest and 8 sites in strips on each of /5 transects D/50 sites totalF.

Scientific name Common name "ody 3ab 3unt *iet rophic !itter $ep $orest Strip

apirus terrestris South American apir 50=.= 8 4 4 / / 46 5 0

anthera onca #aguar E4.> 8 4 4 4 5 40 0 0

uma concolor uma 87 6 4 7 4 5.E 5= 8 /

Catagonus 1agneri Chacoan eccary 48.6 5 4 4 / 5.= 40 4 0

Mazama americana ed brocet *eer 44./ 4 4 5 / / /8 / 0Myrmecophaga tridactyla %iant Anteater 40.> 4 5 / 4 / 4> /5 5

ayassu pecari hite-lipped eccary 5E.> 4 4 8 / / /E 0 0

riodontes ma@imus %iant Armadillo 5=.= 4 4 5 5 /.8 /0.8 0 0

ecari ta9acu Collared eccary 57.> 8 4 8 / 5 /E > 4

Mazama gouazoubira %ray "rocet *eer 50.7 = 4 5 / / /4.8 50 /=

!eopardus pardalis Gcelot =.E 7 / 7 4 /.= 54 / 0

rocyon cancri&orus Crab-eating accoon =.4 8 / 6 5 4.7 /E / /

Cerdocyon thous Crab-eating $o@ 8.5 = 5 6 5 7 // 8 =

(ira barbara ayra 8./ 6 / 8 5 5 50 / 0

!ycalope@ gymnocercus ampas $o@ 7.> = 5 6 5 7 /5 // /6

Nasua nasua South American Coati 7.6 6 / 8 5 4.= 57 / /

(uphractus se@cinctus Si@-banded Armadillo 7.7 4 4 8 5 /.8 > 5 /

3erpailurus yagouaroundi #aguarundi 7.4 7 5 8 4 5.8 /E 0 0

!agostomus ma@imus lains izcacha 7.5 4 4 5 / /.> // 0 0

*asypus no&emcinctus Nine-banded Armadillo 7 E 4 7 5 7 57 0 4

!eopardus geoffroyi %eoffroyPs Cat 4.7 6 5 8 4 5 5/ = //

*asyprocta punctata Central American Agouti 4 4 / 4 / /.4 /6 / /

Conepatus chinga MolinaPs 3og-nosed Sun 5.4 6 / 6 5 4.8 /0.8 // /0

Chaetophractus &illosus !arge 3airy Armadillo 5.4 8 4 4 5 /.6 > 7 5

*olichotis salinicola Chacoan Ca&y 5./ 7 5 4 / /.8 5.E E 8

%alictis cu9a !esser %rison /.> 7 / 7 4 4 50 / 5

olypeutes matacus Southern hree-banded Armadillo /.5 4 4 7 5 / > E 4

*idelphis albi&entris hite-eared Gpossum / E 5 6 5 6.> /0 / 5

Chaetophractus &ellerosus Screaming 3airy Armadillo / 7 4 8 5 5 > / /

Syl&ilagus brasiliensis apeti / 7 4 4 / /.5 57 0 0

a

!ist includes species that either 1ere present at our site or should occur there based on published literature and inter&ie1s.

b ?ey to abbre&iations< "ody, adult body mass DgFJ 3ab, habitat breadth Dlarger &alues indicate more habitat categoriesFJ 3unt, hunting pressure Dlarger &alues sho1 larger hunting pressureFJ *iet,

diet breath Dlarger &alues indicate more diet categoriesFJ rophic, trophic le&elJ !itter, litter size Dnumber of young produced per litterFJ $ep, age at


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analysis to e@amine &ariation in site occurrence as a function of

species ecological traits DAndersen et al., 5000F. e used percent

correctly classified as a measure of predicti&e success DAndersen et al.,

5000F and CohenPs appa statistic to determine o&erall significance of

the optimal classification tree D pacage UUirrPPJ %amer et al., 50/0F.

Second, for species that 1ere modeled for occupancy DN R >F, 1e

conducted a series of 1eighted %!Ms bet1een measures of speciesP

responses to distance along the transect from forested interior Di.e.,

estimates of distance parameter from indi&idual-le&el occupancy

modelsF and all ecological traits of species. he 1eight of each pointin our analysis 1as the in&erse of the standard de&iation of the

distance parameter D"ender et al., />>EF. e fitted models in program

using UUglmPP function and UUidentityPP lin. Models 1ere selected

using Aaie :nformation Criterion 1ith small sample size correction

DA:CcF.

4. esults

4./. Species detection

e recorded 54 species of mammals in our study area, but most

species 1ere detected in less than /0 sampling units Dable /F. Si@

species that should occur in our study area Dbased on published

literature and inter&ie1sF 1ere not detected D9aguar, 1hite-lipped

peccary, giant armadillo, 9aguarundi, plains &izcacha, tapetiF. $i&e

species 1ere found only in continuous forest DSouth American

tapir, Chacoan peccary, red brocet deer, ocelot, tayraF. (ighteen

species 1ere recorded in continuous forest and strips, and the nine-

banded armadillo 1as found e@clusi&ely in forest strips. Modeled

occupancy estimates for the > species recorded in at least /0

sampling units ranged from 0./> DChacoan ca&yF to 0.8= Dgray

brocet deerF Dable A4F.

4.5. elationship bet1een species occurrence and distance along

transects

Species modeled for occupancy &aried in use of continuous

forest and forest strips D$ig. 5, A4F. Gccurrence of giant anteater

and collared peccary decreased near the forest boundary D$ig. 5a

and bF. Gccurrence of gray brocet deer and three-banded

armadillo also appeared to decline gradually along the transect

from forest into strips D$ig. 5c and dF. 3o1e&er, credible inter&als

for the distance parameter 1ere 1ide and o&erlapped zero e@cept in

the case of giant anteater and collared peccary, the t1o species

most clearly associated 1ith forest interior. Gccurrence of pampas

fo@ increased 1ith distance from forest into the strip D $ig. 5fF. 1o

other species appeared to e@hibit similar trends D$ig. 5h and iF, but

detection rates 1ere lo1er Dable /F and credible inter&als for the

distance parameter o&erlapped zero.

4.4. elationship bet1een site of occurrence and species traits

"ody size and habitat breadth 1ere the species traits most

closely associated 1ith site occupancy. A classification tree 1ith

one split based on body size best fit the data and di&ided species

into the follo1ing groups< D/F not detected in study area or in forest

only, and D5F detected in forest and strips. Species 1ith body size

O58 g DN R EF 1ere not recorded or only detected in continuous

forest, 1ith the e@ception of pumas and giant anteaters Dable /F.

hese t1o species 1ere detected in forest sampling units more

often than in strips. Gf the 55 species 1ith body size Q58 g

recorded in the study area, /= occurred in strips, and all these

species e@cept the nine-banded armadillo also 1ere found in forest

Dable /F. his tree correctly classified =EB of the cases, 1hich

1as more than e@pected by chance DCohenPs appa R 0.7E, z R

5.6=, p Q 0.00/F. $or the > species modeled for occupancy,

probability of species occurrence 1ith distance Di.e., from forest

interior and along the forest stripF 1as in&ersely related to body

size and positi&ely related to habitat breadth Dbody size, b R 0.45,

S( R 0.06, t R 8.8/, p Q 0.0/J habitat breadth, b R 0.>7, S( R 0.57, t

R 4.E7, p Q 0.0/J best fit %!M model, $ig. 4, able A7. Although

local informants indicated that most of mammal species in our

study 1ere hunted, this &ariable, along 1ith diet breadth, trophic

le&el, and litter size, did not emerge in models as important for

understanding the relationship bet1een site of occurrence andspecies traits Dable A7F.

7. *iscussion

7./. %eneral patterns of occurrence


6/10


Almost =0B of species cited in the literature for our study area

1ere either not detected or occurred in less than /0 sampling units,

including all species categorized by :CN D50/8F as threatened

D1hite-lipped peccary, &ulnerableJ Chacoan peccary, endangeredJ giant

armadillo, &ulnerableJ and South American tapir, &ulnerableF, 1ith one

e@ception. he giant anteater D&ulnerableF occupied O/0B of the sites

but 1as significantly more liely to occupy continuous forest sites.

!o1 rate of occurrence of mammals in both forest and strips could be

e@plained because medium to large mammals naturally occur at &ery

lo1 densities in our study area and thus 1ere not detected 1ith our sampling effort Di.e., a sampling effectF, or because this fauna has

suffered from human impacts e&en in continuous forest. Another

potential e@planation is that cameras 1ere not effecti&e in detecting

mammals, but this is unliely because trigger speed and sensiti&ity of

cameras 1ere ade)uate for our species, and 1e follo1ed camera-

trapping protocols of other studies of these species De.g., Maffei et al.,

5007J

Body size (kg)

$ig. 4. elationship bet1een body size and habitat breadth and the distance parameter

from occupancy models, as determined by the 1eighted regression analysis. he

shaded areas represent >8B confidence inter&als.

hornton et al., 50//F. $or some large-bodied species that naturally

occur at lo1 densities throughout their range De.g., 9aguar and giant

armadilloF, lac of detection may be related to sampling effect.

3o1e&er, the puma, a large-bodied, 1ide-ranging species that also

occurs at lo1 densities throughout its range Dmost of North and South

AmericaF, 1as detected in 6 sampling units. $or many mammals that

Chaco shares 1ith tropical forest in Central and South America, Chaco

forest Dparticularly in Argentina and araguayF corresponds to the

southernmost part of their distributions De.g., 9aguar, ocelot, peccaries,

South American tapirJ :CN, 50/8F. At limits of species distributions,

optimal en&ironmental conditions and resources may occur 1ith less

fre)uency than in the center of their distributions DCase et al., 5008F,resulting in lo1er population densities and, thus, potentially limiting

$ig. 5. elationship bet1een distance along the transect and occupancy of mammals as determined from hierarchical occupancy models. *istance &alues correspond to distance DmF from the forest

edge D0F 1ith negati&e &alues for continuous forest Dgray areaF and positi&e &alues for forest strips D1hite areaF. >8B "ayesian credible inter&als for parameter estimates of distance co&ariate Db distF are

sho1n in parentheses. *ashed lines correspond to models 1ere credible inter&als for b disto&erlapped zero.


7/10

Dis

tance !ar aeter

*abitat breath

Distance !ar aeter


detection in our study area. 3o1e&er, in protected areas in "oli&ian

Chaco, large mammal densities are e)ual to or larger than in many

tropical forests to the north DMaffei et al., 5007F. hus, if lo1 densities

of mammals are related to geographic boundaries of distributions, this

phenomenon is liely limited to southernmost Chaco and not the entire

Chaco region. Although the lo1 occurrence that 1e documented for

many species may be partially e@plained by sampling effect or

geography, this lo1 occurrence also liely relates to human impacts on

Argentine Chaco. hroughout the region, forest has been highly

modified and degraded as a conse)uence of grazing and logging, andhunting is se&ere though poorly documented for most species

DAltrichter, 5008F.

7.5. se of forest strips

Although the mammal species most &ulnerable to con&ersion of

continuous forest to forest strips already may ha&e been lost from our

study area, /7 of the 57 species remaining 1ere absent from strips or

1ere recorded most fre)uently in continuous forest. :n contrast, only 4

medium-sized carni&ores and possibly the ninebanded armadillo may

benefit from con&ersion of forest to a landscape characterized by

forest strips around agricultural fields based on our occurrence data.

Gur study is consistent 1ith other research that documents lo1er use

of strips than continuous forest by dung-beetles, birds and arborealmammals D"arlo1 et al., 50/0J 3a1es et al., 500EJ !aurance and

!aurance, />>>J !ees and eres, 500EJ ehling and *iemann, 500>F.

3o1e&er, in contrast to our results, medium to large mammals in the

Amazon ha&e been reported to use strips more than continuous forest

D"arlo1 et al., 50/0F. Some mammals 1ere shared bet1een the

Amazon study and our study De.g., collared peccary, giant armadillo,

and othersF, and strip 1idths 1ere comparable D/00 mF. *ifferences in

results of the t1o studies may relate to the landscape matri@

surrounding forest strips, 1hich comprised (ucalyptus plantations in

the Amazonian study and pasture or soybeans in Chaco. $orest

mammals foraged in understory of (ucalyptus D"arlo1 et al., 50/0F.

!ocal lando1ners at our study site reported seeing mammals in strips

or at the edge of strips, but rarely in the agricultural matri@. Soybean

and pasture matri@ of Chaco may pro&ide less suitable habitat and may be less permeable to mo&ement of forest species, as obser&ed for birds

in Chaco forest DMastrangelo and %a&ing, 50/7F. Alternati&ely,

discrepancies bet1een the t1o studies could be related to hunting

pressure if hunting of mammals is higher in strips as compared to

forest in Chaco, and the re&erse occurs in Amazonia. 3unting taes

place at both sites D"arlo1 et al., 50/0F, but relati&e hunting pressure

in forest &ersus strips is unno1n.

7.4. atterns of occurrence and species traits

Studies of forest fragmentation ha&e demonstrated that forest

specialists often are more impacted by forest loss and

fragmentation than generalist species particularly because of

insufficient resources and sensiti&ity to edge effects D3enle et al.,5007J eres, 500/F. !arge-bodied species that range 1idely De.g.,

large carni&oresF often disappear from fragments because their area

re)uirements are not met, and also forest fragmentation increases

their &ulnerability to other threats such as hunting. Although large

blocs of continuous forest remain in our study area, for some

1ide-ranging species such as 1hite-lipped peccaries and 9aguars,

this forest already may be too fragmented to sustain populations

DVuiroga et al., 50/4F. !arge-bodied species and species that

specialize on forest habitat either 1ere not recorded in our study

area or occupied forest sites more often than strips. his group

included herbi&ores Dbro1sersLgrazers and frugi&oresF, omni&ores,

myrmecophages, and carni&ores. All species recorded more

fre)uently in strips than in continuous forest had body mass less

than /0 g and 1ere habitat generalists, occupying a &ariety of open habitats De.g., grasslands and scrublandsF as 1ell as forest

across their ranges DMares et al., />E>J Cane&ari and accaro,

500=J allace et al., 50/0F.

he degree to 1hich occupancy of mammals in our study area

is influenced by )uantity &ersus )uality of habitat, or other factors

such as &ulnerability to hunting, is unno1n, but multiple factors

liely are important. :n this region, con&ersion of forest to

agricultural fields 1ith forest strips results in immediate loss of

=EB of forest area Ddepending on local regulationsJ %inzburg et al.,

50/5F, drastically reducing habitat for forest specialists. Wet,

understanding of habitat )uality for mammals in Chaco forest andstrips is limited. esearch on &egetation has focused primarily on

forest structure. $orest strips, as 1ell as edges of continuous forest,

ha&e higher density of shrubs and small trees and lo1er abundance

of large trees than in forest interior D%inzburg et al., 50/5F. hese

habitat changes should produce more bro1se, and possibly higher

)uality bro1se, in strips for species such as tapirs and deer.

3o1e&er, 1e detected tapirs and red brocet deer only in

continuous forest, and occupancy of grey brocet deer 1as similar

or slightly lo1er in strips than forest. Many mammals in this forest

eat seeds and fruits Dallace et al., 50/0F, and 88B of the 1oody

flora of Chaco is dependent on mammalian dispersal Deriago et al.,

50/8F. :f mammal density is lo1 in strips, either because important

food trees are less abundant in strips or for other reasons, this could

reduce recruitment of food trees leading to further negati&efeedbac on mammal populations. *uring our study, zoochoric

fruits appeared to be less abundant in the forest strips than in forest

interior, but more detailed studies are needed to understand lins

bet1een con&ersion of forest to strips and habitat )uality for

mammals Deriago et al., 50/8F.

rior studies conducted on medium-large mammals in

Argentine Chaco point to hunting as a significant factor in

population declines DAltrichter, 5008J Vuiroga et al., 50/4F. hus,

lac of statistical support for hunting as a predictor of species

occurrence in our study 1as une@pected. his may ha&e occurred

for se&eral reasons. $irst, 1ith only three categories, our inde@ of

hunting based on inter&ie1s may ha&e been too simple to fully

represent differences in hunting pressure among species. Second, a

significant part of the hunting pressure may come from huntersli&ing in urban areas, 1hich is &ery difficult to )uantify and not

included in our models. hird, and perhaps most important, many

species 1ere detected by fe1 or no cameras, precluding analyses of

occupancy. hese species may be locally e@tinct or at lo1 densities

as a result of hunting, or because of combined effects of hunting,

forest loss and fragmentation. 3unting is a ma9or dri&er of

occupancy patterns for mammals in fragmented tropical forest, and

body size and &ulnerability to hunting often are positi&ely

correlated Dhornton et al., 50//J eres, 500/F. :n our study area

many small-bodied species also are hea&ily hunted, potentially

because larger game species ha&e been depleted, but also because

some of these species such as three-banded armadillo are highly

preferred by hunters DAltrichter, 5006F. G&erall, ==B of the

mammals that occurred, or should occur, in our study area arehuntedJ 84B of these species are hea&ily hunted in our study area

and throughout Chaco, including all large-bodied species DO58 gF

e@cept the anteater DAltrichter, 5008, 5006J Vuiroga et al., 50/4F.

he only hea&ily hunted species that commonly occurs in strips and

continuous forest in our study area is the gray brocet deer, 1hich

is a habitat generalist that uses agricultural areas as 1ell as forest

Dallace et al., 50/0F.

7.7. Conser&ation implications

%i&en that strips of forest are mandated by la1 in many tropical

and subtropical areas and 1idely accepted by large lando1ners

D3a1es et al., 500EJ Seghezzo et al., 50//F, forest strips liely 1ill

comprise an increasingly large portion of remaining forest, not only

in Chaco, but also in other regions undergoing rapid forest


8/10


con&ersion for agriculture D$AG, 50/5F. (&idence from our studies

and others D!ees and eres, 500EF demonstrates that these strips

contain only a subset of the original fauna in these forests. :n the

case of Chaco, as forest is con&erted to forest strips embedded in

agricultural fields, loss of forest specialists and persistence of

generalists lead to functional homogenization of biodi&ersity,

1hich may be accompanied by loss of &ital ecosystem ser&ices

such as seed dispersal by forest mammals D*e&ictor et al., 500EJ

uechagut et al., 50/4F. Gur study liely underestimates long-term

effects of forest transformation. Strips in our study area arerelati&ely recent Di.e., Q/0 years oldF and undoubtedly harbor

e@tinction debts Dilman et al., />>7F. As agriculture continues to

e@pand in Chaco, alternati&e landscape configurations urgently

need to be designed, e&aluated, and incorporated into policy if the

mammalian fauna of Chaco is to be retained. Ample e&idence

points to the importance of large fragments and aggregated habitat

to pre&ent local e@tinctions of 1ildlife Dybici and 3ansi, 50/4F.

etention of forest strips may ha&e detrimental impacts on

agricultural production because, for e@ample, trees shade crops and

compete 1ith crop plants for 1ater D%inzburg et al., 50/5F.

Conser&ing the same total forest area that is destined for strips, but

in large interconnected blocs, may recei&e support from pri&ate

lando1ners, 1ho control most of the forested areas in northern

Argentina. :n our study area, lando1ners ha&e sho1n interest inincreasing biodi&ersity &alue of their land, and a conser&ation

payment program associated 1ith the 500> Argentine National

$orestry !a1 pro&ides an opportunity to e@plore, and potentially

finance, multi-sectorial conser&ationLproduction schemes.

An important point that emerges from our study is the critical

situation of the fauna of Chaco. his fauna is poorly studied, di&erse,

and sub9ect to multiple interacting threats. *i&ersity of medium and

large terrestrial mammals of Chaco is comparable to tropical forests

such as the Amazon DMares, />>5J edford et al., />>0F. Armadillos

reach their pea di&ersity in Argentine Chaco D2uleta and "olo&ic,

/>>7F. he endemic Chacoan peccary 1as no1n only from fossil

records until />=7 Detzel et al., />=8F. he fe1 studies of medium to

large mammals that ha&e been conducted in this region primarily focus

on 9aguars and hunting of peccaries DAltrichter, 5008, 5006J Vuiroga etal., 50/4F. hese studies conclude that hunting of Chacoan peccaries

and 1hite-lipped peccaries is not sustainable and that local e@tinction

of 9aguars may be imminent in Argentine Chaco D Altrichter, 5008J

Vuiroga et al., 50/4F. Sustainable har&est management strategies ha&e

been 1idely implemented in tropical regions of South America De.g.,

"odmer et al., />EEF, but are lacing, and particularly challenging, in

Chaco DAltrichter, 500EF. Conser&ation challenges of Chaco 1ere

captured more than t1o decades ago by edford et al. D/>>0F, 1ho

referred to Argentine Chaco forest as UUone of the greatest, yet least

no1n, ecological catastrophes in South AmericaPP. he 500>

Argentine National $orestry !a1, 1hich re)uires that all pro&inces

produce land use plans to conser&e forest DSeghezzo et al., 50//F, pro&ides an unprecedented opportunity to address these challenges and

design a more functional forest-agriculture landscape. 3o1e&er, recent

analyses pro9ect that forest area in Chaco 1ill decline substantially

under the current implementation plan for this la1, and that

ecoregional planning across Chaco is re)uired to maintain forest co&er

and connecti&ity Di)uer-odriguez et al., 50/8F. :n addition to

designing alternati&e landscapes, synergisms bet1een the

configuration of the Chaco landscape and threats such has hunting

need to be addressed as part of the planning process if the UUempty

forestPP syndrome is to be a&oided DAltrichter, 5008J edford, />>5J

eriago et al., 50/8F.

Acno1ledgements

e than S. Andrade, N. Cruz, . %uanuco, . !opez, C. rigo, C.

rucco, . once, #. $einsinger, and Nunez %, C., for help in the field

and &aluable comments. e also than Ministry of (n&ironment of

Salta and Sociedad ural de Salta for logistic support. e than

obert *orazio for assistance 1ith "ayesian models. e are grateful

to *. umiz, and . "ar)uez for insights on species traits. e than .

#ansen and an anonymous re&ie1er for impro&ements on earlier

&ersion of this 1or. his pro9ect 1as funded by Scott Neotropical

$und, ufford Small %rants, $Ps ropical Conser&ation and

*e&elopment rogram, and the Nunezegueiro family.

Appendi@ A. Supplementary material

Supplementary data associated 1ith this article can be found, in the

online &ersion, at http


9/10

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