Ann. Zool. Fennici 45: 281–285 ISSN 0003-455X (print), ISSN 1797-2450 (online) Helsinki 27 August 2008 © Finnish Zoological and Botanical Publishing Board 2008

The small of Warsaw as inferred from tawny ( aluco) pellet analyses

Jakub Gryz, Dagny Krauze & Jacek GoszczyÉski

Division of Forest Zoology and Wildlife Management, Forest Faculty, University of Life Sciences, Nowoursynowska 159, PL-02-776 Warsaw, Poland (corresponding author’s e-mail: jakub.gryz@ wl.sggw.waw.pl)

Received 16 Mar. 2007, revised version received 16 Oct. 2007, accepted 17 Oct. 2007

Gryz, J., Krauze, D. & GoszczyÉski, J. 2008: The small mammals of Warsaw as inferred from (Strix aluco) pellet analyses. — Ann. Zool. Fennici 45: 281–285.

In order to estimate small diversity in Warsaw we analysed pellets of tawny inhabiting green areas. Altogether we found 21 of mammals (9 species in the city centre and 19 on the outskirts). In the central zone the largest share was com- SULVHGRIVWULSHGÀHOGPRXVHKRXVHPRXVHDQGUDWVZKLOHVSHFLHVFRQQHFWHGWRIRUHVW and open areas dominated in the outskirts.

Introduction   ,WV GLHW WKXV YDULHV UHÁHFWLQJ WKH DYDLO- DELOLW\RISUH\ 0LNNROD :LOOLV DVZHOO Where technical problems and the intensive as the species composition of small and medium- human use of urban green space leave traditional sized mammals present in a given area. methods of detecting small mammals (like traps 7KHWDZQ\RZOLVDQRQPLJUDWRU\WHUULWRULDO DQG QHWV  GLIÀFXOW WR DSSO\ DQDO\VHV RI RZO species. Territory sizes relate to the abundance of pellets can offer an alternative source of data SUH\ZLWKWKLVEHLQJPHGLDWHGE\KDELWDWIHUWLOLW\ on the abundances and local distributions of and landscape structure. While radio tracking VPDOOPDPPDOV

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central and outlying areas of Warsaw by verify- DV DQ LWHP RI QDWLRQDO KHULWDJH )XUWKHUPRUH ing a hypothesis that in line with the increasing the relative constancy to the distribution of owl urbanization within the city limits: territories allowed for the collection of pellets in YLUWXDOO\WKHVDPHSODFHVDVLQWKHV 1. the share of insectivores and species con- We analysed pellets collected by following a nected with forest and open areas will VWDQGDUG SURFHGXUH LQYROYLQJ WKH LGHQWLÀFDWLRQ GHFUHDVH of prey species consumed on the basis of the 2. the share of synantropic species as well as VNXOOVIRXQG/DERUDWRU\DQDO\VHVRIWKHPDWHULDO RIWKHVWULSHGÀHOGPRXVHApodemus agrar- PDLQO\UHOLHGXSRQWKHNH\IURP3XFHN   ius, regarded by some authors as a synurbic DQGRWKHUSDSHUV 5XSUHFKW:ROIet species (i.e. Andrzejewski et al   will al.  %DOĀLDXVNLHQē et al   +RZHYHU LQFUHDVHDQG reference was also made to specimens from  WKH GLYHUVLW\ RI WKH VPDOO PDPPDO DVVHP- skulls held by the Division of Forest Zoology blage will decline towards the city centre. and Wildlife Management of the Warsaw Uni- YHUVLW\RI/LIH6FLHQFHV We were further able to compare the results $OWRJHWKHU ZH FROOHFWHG  SHOOHWV IURP of these studies with data obtained from the ZKLFKWKHUHPDLQVRIPDPPDOVZHUHLGHQWL- V *RV]F]\ĸVNLet al-*RV]F]\ĸVNL ÀHG IURPWKHFHQWUDO]RQHDQGIURPWKH XQSXEO GDWD  WR VHH ZKHWKHU RQJRLQJ XUEDQL- RXWVNLUWV  ,Q WRWDO  PDPPDOV  DQG  zation of the city has affected small-mammal IURPWKHFHQWUDO]RQHDQGWKHRXWVNLUWVUHVSHF- assemblages. WLYHO\ ZHUHLGHQWLÀHGWRVSHFLHVOHYHO0DPPDOV IRU ZKLFK LGHQWLÀFDWLRQ WR VSHFLHV OHYHO SURYHG impossible were not included in calculations of Study area and methods VSHFLHVGLYHUVLW\+RZHYHUSUH\LWHPVLGHQWLÀHG to the level of the order or (Insectivora :HFRQGXFWHGRXUVWXG\LQWKHFDSLWDORI3RODQG Microtus or Rattus ZHUHXVHGWRWHVWWKHÀUVWDQG EHWZHHQ 1RYHPEHU  DQG 2FWREHU  the second hypotheses. :DUVDZFRYHUVNP2 DQGKDVPRUHWKDQ 7KH 6KDQQRQ:LHQHU GLYHUVLW\ LQGH[ ZDV million inhabitants. Built-up areas account for FDOFXODWHGXVLQJEDVH .UHEV   RI WKH DUHD DUDEOH ODQG DQG RUFKDUGV IRU 6WDWLVWLFDO DQDO\VHV LQFOXGHG D WHVW IRU WKH IRUHVWVIRUDQGZDWHUVIRURI FRPSDULVRQ RI WZR SHUFHQWDJHV %DLOH\   the total area. DQGWKH0DQQ:KLWQH\:LOFR[RQWHVW To estimate the composition of small- PDPPDODVVHPEODJHVZHVHOHFWHGWHUULWRULHV RIWDZQ\RZOVIURPZKLFKSHOOHWVZHUHFROOHFWHG Results and discussion DWOHDVWRQFHDPRQWKRYHUD\HDUSHULRG(LJKW of the territories were situated in the central zone $OWRJHWKHUWKHUHPDLQVRIVSHFLHVUHSUHVHQW- LQ GLVWULFWV ZLWK PRVW LQKDELWDQWV  DQG HOHYHQ ing four orders were collected: RodentiaInsec- on the outskirts of Warsaw. Tawny owl territories tivoraChiroptera and Carnivora (Table 1). The ZHUHORFDWHGLQPXQLFLSDOSDUNVFHPHWHULHVDQG QXPEHURIPDPPDOVSHFLHVLGHQWLÀHGZDVWZLFH woodlands. as high in the outskirts of Warsaw as compared To indicate changes in the small mammal ZLWKWKDWLQWKHFHQWUDO]RQH DQGVSHFLHV IDXQDWKDWPLJKWKDYHRFFXUUHGVLQFHWKHV respectively). Rodents were caught most often ZHVHOHFWHGRQHVLWHIURPZKLFKVXIÀFLHQWQXP- DFFRXQWLQJIRURIWKHSUH\LWHPV ZLWKWKH bers of pellets were collected in both study greatest shares being accounted for by the striped periods to allow for statistical comparisons. It ÀHOGPRXVHDQGWKH\HOORZQHFNHGPRXVHApo- appeared that the most suitable site for this GHPXVÁDYLFROOLV. ZDVâD]LHQNL3DUNZKLFKLVORFDWHGLQWKHFLW\ The composition of the small-mammal FHQWUH,WVDUHDRIFDKDKDVQRWEHHQVXEMHFW assemblages in the central and outlying parts WR DQ\ PDMRU FKDQJHV DV LW KDV EHHQ SURWHFWHG of Warsaw differed markedly (Table 2). In line

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ZLWK RXU DVVXPSWLRQV WKH VWULSHG ÀHOG PRXVH Table 1. Mammals in the diet of the tawny owl in was caught more often in the central zone. As far Warsaw. DV V\QDQWKURSLF VSHFLHV DUH FRQFHUQHG ZH FRQ- Prey Species Species present ÀUPHGWKDWUDWVRattus spp. were more numerous number LQWKHFHQWUDO]RQH+RZHYHULQWKHFDVHRIWKH Central Outskirts house mouse Mus musculus — also regarded as zone

V\QDWKURSLF³WKHGLIIHUHQFHZDVQRWVLJQLÀFDQW Apodemus agrarius 89 42 Neither forest species like the yellow-necked Apodemus flavicollis – 105 mouse and the bank vole Myodes glareolusQRU Apodemus spp. 12 27 species of open areas like the common vole Mus musculus 67 Microtus arvalisWKHURRWYROHM. oeconomus and Micromys minutus 37 M. agrestis were collected from the Rattus norvegicus 810 WKHÀHOGYROH Rattus spp. 5 – central zone. Insectivores were only represented Microtus arvalis –29 by the mole Talpa europaeaDOWRJHWKHUVXSSRUWHG Microtus oeconomus –8 RXUÀUVWK\SRWKHVLV 7DEOH 6LPLODUGLIIHUHQFHV Microtus subterraneus 25 9 ZLWK WKH H[FHSWLRQ RI Rattus VSS ZKHUH WKHUH Microtus spp. – 10 Arvicola terrestris –1 ZDV QR VLJQLÀFDQW GLIIHUHQFH  EHWZHHQ FHQWUDO Myodes glareolus –63 zone and the outskirts were found when we com- Sciurus vulgaris 1– pared mean percentage shares from separate terri- I Rodentia 11 149 318 WRULHVE\PHDQVRIWKH0DQQ:KLWQH\:LOFR[RQ Talpa europaea 725 WHVW6SHFLHVGLYHUVLW\FDOFXODWHGXVLQJWKH6KDQ- Sorex araneus –28 Sorex minutus –2 QRQ:LHQHU LQGH[ ZDV KLJKHU LQ WKH RXWVNLUWV Sorex spp. – 3  WKDQLQWKHFHQWUDO]RQH   Erinaceus concolor –1 6LJQLÀFDQW FKDQJHV LQ DEXQGDQFH DQG UDQJH Insectivora indet. 3 – VL]HLQFRPSDULVRQZLWKWKHVZHUHIRXQGLQ I Insectivora 4 10 59 WKHVWULSHGÀHOGPRXVHKRXVHPRXVHPROHDQG Nyctalus noctula 59 Myotis nattereri –6 EDWV 7DEOH 7KHSHUFHQWDJHRIKRXVHPLFHLQ Myotis daubentonii –1 WKH GLHW RI WDZQ\ RZOV LV FXUUHQWO\ VLJQLÀFDQWO\ Eptesicus serotinus 9– ORZHU WKDQ LQ WKH SDVW SUREDEO\ EHFDXVH WKLV I Chiroptera 4 14 16 species continues to decrease in abundance in Mustela nivalis –2 WKHFHQWUDO]RQH/LYHWUDSSLQJVWXGLHVFRQGXFWHG Mustela vison –1 Carnivora 2 0 3 LQ :DUVDZ FRQÀUP WKLV YLHZ $QGU]HMHZVNL et I I Prey 173 396 al.  &LFKRFND   :H DOVR UHJLVWHUHG I Species 21 9 19 DQ LQFUHDVH RI SHUFHQWDJH VKDUH RI ÀHOG VWULSHG

Table 2. Small mammal assemblage on the basis of pellet analyses.

Prey All items Selected territories

Central Outskirts Central zone Outskirts zone (%) (%) (% ± SD) (% ± SD)

Apodemus agrarius 75.4 12.6 p < 0.001 82.5 ± 15.1 14.0 ± 8.4 Apodemus flavicollis 0 31.5 p < 0.001 0 ± 0 28.2 ± 18.9 Mus musculus 5.1 2.1 p > 0.05 3.4 ± 4.2 02.5 ± 2.6 Rattus spp. 11.0 3.0 p < 0.01 7.7 ± 6.8 03.5 ± 5.0 Myodes glareolus 0 18.9 p < 0.001 0 ± 0 17.4 ± 10.3 Microtus spp.* 0 14.1 p < 0.001 0 ± 0 15.8 ± 11.1 Insectivora 8.5 17.3 p < 0.01 6.4 ± 7.3 18.5 ± 5.5 Total mammals 118 333 118 000329

*without Microtus subterraneus.

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Table 3. Small mammal assemblage in Łazienki Park (central zone) in the two periods.

Prey 1975–1977* 2003–2006

Percentage n Percentage n

Apodemus agrarius 27.9 26 48.9 46 p < 0.01 Mus musculus 23.7 22 3.2 3 p < 0.001 Rattus norvegicus 7.5 7 5.3 5 p > 0.05 Microtus subterraneus 19.4 18 26.6 25 p > 0.05 Talpa europaea 21.5 20 1.1 1 p < 0.001 Chiroptera 0 0 14.9 14 p < 0.001 Mammals total 93 94

*GoszczyÉski et al. 1993, J. GoszczyÉski unpubl. data.

mouse in the diet of tawny owl. This is in line References ZLWKWKHODWHVWOLYHWUDSSLQJVWXGLHVIRUP:DUVDZ which showed clear dominance of this species in $QGU]HMHZVNL 5 %DELĸVND:HUND - *OLZLF] -  *RV]F]\ĸVNL-6\QXUEL]DWLRQSURFHVVLQSRSXOD- the small mammal assemblage of city parks in the tion of Apodemus agrarius , &KDUDFWHULVWLFV RI SRSX- FHQWUHRI:DUVDZ &LFKRFND-%DELĸVND lations in urbanization gradient. — Acta Theriol  Werka pers. comm.). We noticed lower numbers ² RIPROHVEHLQJFDXJKWE\RZOVLQâD]LHQNL3DUND %DLOH\17-Statistical methods in biology³&DP- fact which may readily be ascribed to persecution EULGJH8QLY3UHV&DPEULGJH/RQGRQ8. %DOĀLDXVNLHQē/$QDO\VLVRIWDZQ\RZO Strix aluco) (the introduction of new methods of eradicating food remains as a tool for long-term monitoring of small moles and efforts to prevent them from forming mammals. — Acta Zoologica Lituanica² PROHKLOOVRQODZQV DVZHOODVWKHOLNHO\LVRODWLRQ %DOĀLDXVNLHQē/-XåNDLWLV5 0DçHLN\Wē5,GHQ- RIWKHSRSXODWLRQ,Q3RODQGEDWVKDYHWHQGHGWR WLÀFDWLRQ RI VKUHZV DQG URGHQWV IURP VNXOO UHPDLQV LQFUHDVHLQQXPEHUVRYHUWKHODVWIHZGHFDGHVVR according to the length of a tooth row. — Acta Zoologica Lituanica² a greater share of this group in the diet of tawny &LFKRFND$  Small mammals of the green areas of RZOVLVLQOLQHZLWKWKHJHQHUDOWUHQG /HVLĸVNLet Warsaw ³06FWKHVLV:DUVDZ8QLYHUVLW\RI/LIH6FL- al  HQFHV>,Q3ROLVKZLWK(QJOLVKVXPPDU\@ As we are aware that seasonal and local dif- &ODUN'5 %XQFN&07UHQGVLQ1RUWK$PHULFDQ ferences in the prey composition of the tawny owl small mammals found in common pellets ( alba) dietary studies. — Can. J. Zool. ² FRXOGLQÁXHQFHRXUUHVXOWVWRPDNHWKHPDWHULDO *RV]F]\ĸVNL -  7KH HIIHFW RI SUHGDWRU\ ELUGV DQG comparable we collected pellets seasonally and at mammals on populations of their prey. — Wiad. Ekol. the same time in the two zones (in each of them ² in many territories located in diverse habitats). *RV]F]\ĸVNL-&RPSDUDWLYHDQDO\VLVRIIRRGRIRZOV in agrocenoses. — Ekol. pol. 7KHUHIRUHZHEHOLHYHWKDWRXUGDWDVKRZDUHOL- ² *RV]F]\ĸVNL--DEãRĸVNL3/HVLĸVNL* 5RPDQRZVNL- able view of the differences between diet com- 9DULDWLRQLQGLHWRI7DZQ\RZOStrix aluco/DORQJ position of the tawny owl population both in the an urbanization gradient. — Acta Orn.² outskirts and in the city centre. .UHEV & -  Ecology. The experimental analysis of distribution and abundance³+DUSHU&ROOLQV&ROOHJH 3XEOLVKHUV1HZ,Q3ROLVKZLWK(QJOLVKVXPPDU\@ 5yŧ\FNLDQG$GDP7DUãRZVNLIRUWKHLUKHOSLQFROOHFWLQJSHO- /HVLĸVNL**U\]- .RZDOVNL0'RHVWKHGLHWRI OHWV*U]HJRU]/HVLĸVNLDQG-DQ&LFKRFNLKHOSHGXVLGHQWLI\ DQRSSRUWXQLVWLFUDSWRUWKHWDZQ\RZOStrix alucoUHÁHFW SUH\UHPQDQWV:HH[SUHVVRXUJUDWLWXGHWRIUHHDQRQ\PRXV long-term changes in bat abundance? A test in central referees of Annales Zoologici Fennici for their valuable com- Poland. — Folia Zoologica>,QSUHVV@ ments on the previous version of the paper. The language was /RYH 5$ :HEEHQ & *OXH ' (  +DUULV 6  checked by James Richards. &KDQJHVLQIRRGRI%ULWLVKEDUQRZOV Tyto alba) between

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DQG³Mammal Rev² 3ROLVKZLWK(QJOLVKVXPPDU\@ /XQLDN 0  ,QYHQWRU\ RI WKH DYLIDXQD RI :DUVDZ 6XQGH3 %¡OVWDG06$WHOHPHWU\VWXG\RIWKH ³VSHFLHVFRPSRVLWLRQDEXQGDQFHDQGKDELWDWGLVWULEX- social organisation of a tawny owl (Strix aluco) popula- tion. — Acta Orn² tion. — J. Zool., Lond² 0LNNROD+ :LOOLV,Owls of Europe. — T. & A.D. 6XQGH 3 2YHUVNDXJ . %ROVWDG - 3  ‘LHQ -  3R\VHU/RQGRQ /LYLQJ DW WKH OLPLW HFRORJ\ DQG EHKDYLRXU RI 7DZQ\ 0RUHQR (  %DUERVD $  'LVWULEXWLRQ SDWWHUQV RI 2ZOVStrix aluco in a northern edge population in central small mammal fauna along gradients of latitude and Norway. — Ardea² HOHYDWLRQLQ1RUWKHUQ6SDLQ³Zeitschrift für Sugetier- :DVLOHZVNL-  '\QDPLFV RI WKH DEXQGDQFH DQG FRQ- kunde² VXPSWLRQ RI ELUGV RI SUH\ LQ WKH 1LHSRãRPLFH )RUHVW 3XFHN =  Keys to vertebrates of Poland: mammals. — Acta Zool. Cracov² ³3ROLVK6FLHQWLÀF3XEOLVKHUV:DUVDZ :ROI36WUDVFKLO+ %DXHU.Rattus rattus und 5HGSDWK60+DELWDWIUDJPHQWDWLRQDQGWKHLQGLYLG- Rattus norvegicus in Österreich und deren Unterschei- ual: tawny owls Strix aluco in woodland patches. — J. GXQJDQ6FKlGHOXQGSRVWFUDQLDOHP6NHOHWW³Mitt. Abt. Anim. Ecol² Zool² 5XSUHFKW $ /  &ULWHULD IRU VSHFLHV GHWHUPLQDWLRQ ,Q3ROLVKZLWK(QJOLVKVXPPDU\@ =DOHZVNL$'LHWRIXUEDQDQGVXEXUEDQWDZQ\RZOV 5XSUHFKW$ / $ NH\ IRU PDQGLEOH LGHQWLÀFDWLRQRI (Strix aluco) in the breeding season. — J. Raptor Res. Polish bats. — 3U]HJOćG =RRORJLF]Q\  ² >,Q ²

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