original research article
received: 30.11.2018 r., peer-reviewed: 5.02.2018 r., accepted: 23.08.2018 2018 R.W. Mysłajek et al.
e-ISSN 2082-8926
Composition of the wolf's Canis lupus L. diet in the Wigry National Park
Robert W. Mysłajek1*, Sabina Nowak2, Maciej Romański3, Katarzyna Tołkacz4
1 n ers ty o Warsaw, ac lty o B olo y, nst t te o enet cs and B otec nolo y, Paw sk e o a, 02–10 Warsaw, Poland
2 ssoc at on or at re Wol , wardorzeczka 229, 4– 24 L owa, Poland 3W ry at onal Park, rzywe 82, 1 –402 wałk , Poland
4 n ers ty o Warsaw, ac lty o B olo y, nst t te o oolo y, e art ent o Paras tolo y, M eczn kowa 1, 02–09 Warsaw, Poland el. 48 04 2 228, e a l: ro ert. yslajek . w.ed . l
Abstract. the diet of wolves Canis lupus L. was assessed n t e W ry at onal Park n ort astern Poland, w c o erla s w t t e at ra 2000 s te stoja W erska . e content o a total o 149 scat sa les was collected n 2017 and analysed n order to deter ne d etary co os t on. Wol es r ar ly eed on w ld n lates, w c ake 7 .4 o ood o ass.
es te t e act t at w ld oar Sus scrofa l. and red deer Cervus elaphus l. dominate in the ungulate community in the study area, t e r ary rey s ec es was o ser ed to e roe deer Capreolus capreolus L. w t 9. , w le red deer and w ld oar only const t ted 18.7 and 8. o t e ood o ass, res ect ely. dd t onally, ea er Castor fiber L. was o nd to e an ortant rey (10.9 ) as well and l estock acco nted or 1 .1 o all o ass cons ed. e l estock eaten y wol es also ncl ded carcasses of domestic animals illegally disposed of in the forest. We therefore conclude that decisions on the management of the wol s ood ase w t n rotected areas, s c as nat onal arks or at ra 2000 s tes, s o ld e receded y ntens e local st d es.
Keywords: wol d et, rey select on, w ld n lates, ea er, l estock
1. Introduction
Wolves Canis lupus l. prey mostly on large ungulates (new- so e et al. 201 ) t, re onally, ay s ec al se n ot er ood so rces ( ar ont et al. 200 aler et al. 2018). Moreo er, wolves display seasonal and multiannual dietary shifts depend-
n on t e dyna cs o o lat on n ers o t e r otent al rey ( doro c et al. 200 Mer et al. 2011). ere ore, w en lann n act ons to rotect wol es n nat onal arks ( a - rozy 2015) or in natura 2000 sites (Diserens et al. 2017), the knowled e o not only t e d str t on, n ers and de o-
ra c ara eters o o lat ons t also t e s ec es ora - n attern s necessary ( drzejewsk et al. 2010).
e wol s d et co os t on as een e tens ely st d ed n Poland ( drzejewsk et al. 1992 etana and l ek 199 owak et al. 200 , 2011). e ajor rey are lar e w ld n lates (on an a era e, 8 . o t e ood o ass consumed), including red deer Cervus elaphus L. n t e rst lace, ollowed y roe deer Capreolus capreolus l. and wild oar Sus scrofa L. n so e areas, owe er, ea ers Castor
fiber L. are also ntens ely nted y wol es ( owak et al.
2011 drzejewsk et al. 2012).
e wol o lat on n a t n t e st w orest s one o t e est reco n sed o lat ons n ter s o ood co os - tion. however, the north-western edge of the forest, protected
n t e W ry at onal Park (WP ) and t e stoja W er- ska (PL 200004) at ra 2000 s te, as not een n est at- ed ( ar szk ew cz 2011 drzejewsk et al. 2012). t t e sa e t e, t e area, a nly d e to a lar e s are o lakes, d -
ers cons dera ly ro t e re a n n tracts o t e st w orest w t res ect to ot to o ra y and a tats.
e a o t s work was to deter ne t e wol s d et composition as well as the pattern of prey selection from the ungulate mammal community in the WPn.
2. Study area
e WP ( 4 02 2 04 20 ), des nated n 1989, co ers an area o 1 0.8 k 2 w t a rt er 118.2 k 2 or n ts er zone. o t 94 o t e ark land o er-
la s w t t e s ec al area o conser at on stoja W er- ska co er n 1 0.7 k 2. the area is considered a wetland of international importance under the ramsar convention and s also enco assed y t e s ec al rd rotect on area
P szcza stowska (PLB200002).
e at onal Park s located n t e nort western art o t e st w orest n t e Podlask e o odes . e ark s lar ely a orested area ( ). cots ne Pinus sylvestris L. stands re a l n t e orests (80 ), w lst n s aller orest e anses, t ere do nates e t er orway s r ce Picea abies L., lack alder Alnus glutinosa aertn., rc Betula spp. l.
or ed nc late oak Quercus robur L. ( oz sk 2002). e re a n n ort on o t e ark cons sts o water od es (19 ) and non orest co n t es (18 ). e area re a ns nder t e n ence o cont nental cl ate c aracter sed y lon winters, which usually last from the third decade of novem-
er to t e rst decade o r l. e a era e ann al te era- t re s . , and t e a era e ann al rec tat on s 89
(data or t e years 2002–201 , ro ded y t e W ry nte rated n ron ental Mon tor n tat on n o olewo).
o t 19 s ec es o lar e and ed s zed a als a e een recorded n t e WP , ncl d n o r s ec es o wild hoofed mammals, such as red deer, roe deer, european elk and w ld oar ( a rozy 201 ). Lyn Lynx lynx err. s anot er lar e redatory a al, ne t to wol ( drzejewsk et al. 2002 edz ałkowska et al. 200 ). Mon tor n er-
or ed s n ca era tra s and enet c analyses acked y t e crosatell te arkers ro ded e dence t at the WPn area overlaps with the fragments of territories of t e t ree wol acks (Ro a sk et al. 2018).
3. Materials and methods
the wolf’s diet in the WPn was assessed through the analysis of 149 scats collected in 2017, including 101 scats sa led d r n t e a t n–w nter season ( ro 1 cto er to 15 april) and 48 scats sampled in the spring–summer season ( ro 1 r l to 0 e te er). e scats were at ered across t e ent re nat onal ark ( . 1), w t n t e terr tor es o t ree acks o wol es (Ro a sk et al. 2018). e locat on of every scat sample was positioned using the gPs receiver.
Faeces were placed in paper envelopes, dried out and stored until the time of analysing. Dry scats were weighed and rin- sed on a dense mesh, then dried up again and weighed. the
rey s ec es cons ed y wol es were dent ed ased on t e was ed ood re a ns s c as ones, a r, oo es, claws and teet (Lock e 19 9 oszczy sk 1974). e rey s ec es were dent ed to s ec es and/or en s w t t e a d o den- t cat on des ( z rdz k 197 e rot et al. 1982 P cek 1984 eer nk 1991 e Mar n s and s rea 200 ) as well as o t e a t ors own co arat e ater als. n case o do t, the microscope hair preparations were made.
e ood co os t on was e ressed as (1) ercenta e o proportion of scats containing respective food categories in t e total n er o all t e scats collected and (2) ercenta-
e o o ass o nd d al d et te s n relat on to t e total o ass o ood cons ed y wol es. e o ass was de- ter ned y lt ly n t e dry ass o a en cate ory o
ood re a ns y t e ollow n d est l ty coe c ents: e- d s zed a als, 0 n lates, 118 and lant ater al, 4 ( drzejewska and drzejewsk 1998). e Le ns or la (19 8) was sed to calc late t e readt o ood n c e:
1 B = –––––,
pi2 where
pi is the percentage of proportion of every prey category in t e total o ass o ood cons ed.
e Le ns nde ass es t e al es ro 1 (w en only one food item is consumed) to 3 (when an animal evenly uses all food categories). the prey was grouped into the fol- lowing categories for the purpose of calculations: (1) wild ungulate mammals, (2) domestic animals, (3) small wild mammals.
e ood select ty was calc lated ased on t e aco s formula (1974):
r – p D = –––––––––––,
r + p – 2rp where
r s t e ro ort on o a en rey s ec es n t e total n er o w ld n late a als k lled y wol es and p is the pro- portion of a given prey species in the wild ungulate community.
D nde ass es t e al es ro 1 ( ll re erence) to 1 (complete avoiding). the share of respective ungulate mammal s ec es n t e wol s d et was e al ated ased on t e r re en- cy n scats. cer ds co ld not e dent ed to s ec es, t e con- tr t on o res ect e rey s ec es n scats was e al ated ased on the species proportion found in those samples where it was evaluated. Data on the species composition of wild ungulate mammals in the WPn was received from camera traps recor- dings. in the present study, we used recordings from 54 camera tra s a a la le ro 2 locat ons o er t e ent re nat onal ark area t e deta ls o ca era tra s nstallat on were re orted y Ro a sk et al. (2018). e total ater al e raced 10,2 4 ca era tra s days, collected ro an ary to e te er 2017, which contained 5,119 recordings (2,933 independent events) w t ,149 w ld n late a als re stered.
4. Results
the analysis of scat content revealed that the food niche of wolves in the WPn was relatively narrow (B 1. 7). ey are
specialised in hunting wild ungulate mammals, which consti- t ted 7 .4 o t e ood o ass cons ed. Roe deer was t e
ost re ent rey, and ts ro ort on n t e wol s d et was tw ce as as t at o red deer, 9. and 18.7 , res ec- t ely. W ld oar was t e t rd ost s n cant rey, w lst
ro ean elk was only s orad cally cons ed. Bea ers were an ortant e tra rey n t e wol s d et and ro ded or as c as 10.9 o t e ood o ass cons ed. are was rarely cons ed, w ereas s all rodents and rds were only an n re ent rey. cats o wol es ro t e WP conta ned re a ns o a cow, a oat, s and do s, and t e r co ned contr t on n t e ood o ass was 9. . ny eces o
last c were o nd n one o t e scat sa les ( a le 1).
the wild ungulate mammal community in the WPn eva- luated using camera traps recordings consists of four spe- c es. W ld oar was t e ost re ently recorded (n 2,808
nd d als), t e second ost a ndant was red deer (n =
2,273) and roe deer (n 8 ), w lst ro ean elk was least re ent (n 202) ( a le 2). ro t e a o e n late co - munity, wolves tended to select roe deer, for which the ivlev select on coe c ent was D 0.7 . e re a n n s ec es were k lled to a lesser de ree t an was t e r contr t on to t e rey co n ty, w at was corro orated y t e ne at e
al es o t e select on coe c ent ( a le 2).
5. Discussion
several earlier studies provided evidence that red deer is ost re ently nted y wol es n nort eastern Poland and is a prey of preference in the wild ungulate mammal co n ty ( drzejewsk et al. 1992, 2000, 2002a kar-
a 199 drzejewska et al. 1997). rst analyses o t e wol s d et co os t on n t e st w orest n t e years 2000–200 a e also s own t at red deer s t e rey o re-
Figure 1. rea o W ry at onal Park w t locat ons o col- lected wolf scats
erence ( drzejewsk et al. 2012). owe er, a later rese- arc ade w t t e se o a lar e n er o scats (n 7 ), collected in the years 2005–2009, yielded results indicating t at wol es n t e st w orest c ose to rey on roe deer, w c contr ted to 2 o t e wol s d et ( ar sz- k ew cz 2011). r st d es ro de e dence t at t e a o e trend ay e cont n ed, eca se roe deer was also a ajor component of the wolf’s diet in the WPn in 2017.
Bea ers are a s n cant rey te (10.9 o ood o ass) for wolves in the WPn, and this share in the wolf’s diet was er t an t at o t e wol es l n n t e ent re st w orest. e latter s are ct ated ro 4. to 8 o t e ood o ass n t e years 2000–2009 ( ar szk ew cz 2011 drzejewsk et al. 2012). arl er, ea er was nd cated as a prey item in the wolf’s diet in numerous forests located in the w ole lowland art o Poland, n ot t e east ( drzejewsk et al. 2012) and t e west o t e co ntry ( owak et al. 2011). e
lar est contr t on o ea er rey n t e wol s d et was re- corded n t e Ro ncka orest, t at s, 1 .4 o ood o ass,
ollowed y t e kal ska orest (1 .4 ) and t e nyszyn or- est (24. ) ( drzejewsk et al. 2012). Bea er s lack n only n t e rey o wol es l n n t e ar at ans ( etana, l ek 199 owak et al. 200 drzejewsk et al. 2012).
o est c an als, ncl d n ot l estock (cattle, oats and s) and ets, s c as do s, co r sed 9. o t e ood o ass n ested y wol es n t e WP . e a o e al e s lower t an t at re orted earl er or t e ent re e ense o t e st w orest, w c reac ed ro 1 .7 to 1 . o ood o ass, as was deter ned n t e years 2000–2009 y ar-
szk ew cz (2011) and drzejewsk et al. (2012). e scat analys s does not ena le t e dent cat on o w et er t e rey cons ed y an an al was a nted rey or a carcass o nd.
Wol es are known to eed on carcasses ncl d n dead l estock lle ally d ed n t e orest ( drzejewsk et al. 2002a). al carcasses a e een recorded n t e WP (M. Ro a sk , n l s ed data). L kew se, ar szk ew cz (2011) re orted wolves foraging on carcasses of horses, pigs and cows illegally d s osed o n ar o s arts o t e st w orest. e act that residues of domestic animals were detected in wolves’ scats n w nter, eyond t e nor al raz n season, test es to wol es ora n on carcasses o dead an als ( ar szk ew cz 2011).
the meta-analysis of the wolf’s diet over the entire range o wol occ rrence re ealed t e lack o relat ons etween the dietary diversity and the degree of anthropopressure ( ewso e et al. 201 ). e ncreased se y wol es o do est c an als and waste d s osed y eo le was only o nd n t e case o decreased n ers o t e w ld n late a al o lat ons ( doro c et al. 200 ewso e et al. 201 ). e ood de and or wol es n Poland can e lly sec red eca se o t e ncreas n n ers o a e an-
al o lat ons ncl d n red deer, roe deer and w ld oar (Borow k et al. 201 ). e con cts w t l estock ar n a e een l ted y ntrod c n ar o s da a e re ent on eas res d r n cattle raz n ( owak et al. 200 owak and Mysłajek 200 ).
Table 1. et co os t on o wol es n W ry at onal Park
Food item total
B
red deer Cervus elaphus 18.7 18.1
roe deer Capreolus capreolus 9. .9
Moose Alces alces 0.7 1.3
er ds n dent ed Cervidae 8.1 10.7
W ld oar Sus scrofa 8.3 13.4
Wild ungulates total 75.4 7 .
goat Capra aegagrus hircus 0.5 0.7
cattle Bos sp. 4. 1.3
Domestic pig Sus scrofa domesticus 2.0 1.3 Domestic dog Canis lupus familiaris 2.3 4.7
Domestic animals total 9.5 8.1
european hare Lepus europaeus 4.1 .0
Beaver Castor fiber 10.9 14.8
small rodents 0.1 4.7
small mammals total 15.1 25.5
Birds aves + 0.7
Plant material + 4.7
Plastic + 0.7
er o scats 149
B o ass o cons ed ood k 138.9
ood n c e readt B 1. 7
– s are o scats , B – s are o total o ass cons ed . ( ) on- tr t on to d et 0.0
Table 2. Prey select on y wol es ro t e w ld n late co n ty n W ry at onal Park
species r p D
W ld oar Sus scrofa 1 .7 4 . 0. 14
Moose Alces alces 1.7 3.3 -0.327
red deer Cervus elaphus 2 . 37.0 -0.237 roe deer Capreolus capreolus 55.0 14.1 0.7 r – ract on o a s ec es a on all w ld s ec es k lled y wol es [%], p – fraction of species in the ungulate community [%], D – select ty nde a ter aco s (1974)
this study indicates that wolves in the WPn also prey on do s. o s, as do est cated redators, are known to ne at - ely n ence t e local w ldl e ( ra ze ryz and ryz 2014 W erz owska et al. 201 ). ct ons a ed at red c n t e n - er o do s n orests are la o r ntens e and costly, w lst t e r physical elimination through catching or hunting encounters a s stant al soc al o os t on ( ol es et al. 201 ). at s w y wolves preying on dogs that penetrate the natural environment, and rotected areas n art c lar, s o ld e treated as ro ders o a des ra le ecosyste ser ce (Mysłajek 2014).
e enet c str ct re o wol es s correlated, n t e road geographical scale, with the prevailing environment and the rey ty e, w c can e e la ned n ter s o a natal a tat re erence nd ct on (P lot et al. 200 ar c ael et al. 2007 M s an et al. 2007). owe er, d erences were locally recorded n d etary co os t on etween t e adjacent acks o wolves and even amongst respective individuals (Urton and o son 200 ar ont et al. 2009). an es n t e ood co - os t on o t ese redators ay also e s a ed y c an es n the size of potential prey populations (Darimont and reichmen 2002, Mer et al. 2011). s e as ses t e s n cance o systematic research on the wolf's diet composition, as such stu- dies should provide vital information necessary for managing
ot wol s o lat on and o lat ons o t e r otent al rey.
e latter s o art c lar ortance or t e nat onal arks and at ra 2000 s tes, w ere conser at on lans are de elo ed y tak n nto acco nt t e ss e o wol es, and, n arallel, nt n or reduction of wild ungulates is done.
Conflict of interests
e a t ors declare no co et n con cts o nterests.
Acknowledgment and source of funding
s work was nded y t e orest nd o t e tate orests at onal orest old n nder t e roject: ro-
n t e o ser at on syste o lar e and ed a als n orest ecosyste s o t e W ry at onal Park – ta e .
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Authors’ contribution
R.W.M., . . – des ned t e art cle conce t, ela orated t e res lts and wrote t e a er M.R. – ade eld work
. .– er or ed la oratory work.
ranslated y: adw a enk ew cz
