Ann. Zool. Fennici 45: 308–316 ISSN 0003-455X (print), ISSN 1797-2450 (online) Helsinki 27 August 2008 © Finnish Zoological and Botanical Publishing Board 2008

Variation of assemblages in fox and marten faeces collected in southern Poland

Tomasz Skalski1 & Izabela Wierzbowska2

1) Institute of Zoology, Jagiellonian University, Ingardena 6, PL-30-060 Krakow, Poland (e-mail: [email protected]) 2) Institute of Environmental Sciences, Jagiellonian University, Gronostajowa 7, PL-30-387 Krakow, Poland

Received 1 Mar. 2007, revised version received 23 Oct. 2007, accepted 23 Oct. 2007

Skalski, T. & Wierzbowska, I. 2008: Variation of insect assemblages in fox and marten faeces col- lected in southern Poland. — Ann. Zool. Fennici 45: 308–316.

Remains of invertebrates, especially , are frequently found in carnivores’ faeces. Invertebrates are generally restricted to a given area and many factors such as land- use pattern, vegetation structure or even moisture can separate different groups; thus, LQYHUWHEUDWHVFDQEHXVHGDVELRLQGLFDWRUV)RUW\ÀYHVDPSOHVRIPDUWHQDQGIR[VFDWV were analysed for the presence of insect species. Thirty insect species — which were H[SHFWHGWREHIRXQG³ZHUHLGHQWLÀHG&DQRQLFDOFRUUHVSRQGHQFHDQDO\VLVLQGLFDWHG WKDWWKHIDFWRU¶VSHFLHV· LHPDUWHQDQGIR[ ZDVWKHPDLQIDFWRUGHWHUPLQLQJLQYHU- tebrate species variation. Redundancy analysis allowed us to identify invertebrate ecological groups associated with these two carnivores. Martens prefer nest-building insects as a supplementary source of food and mainly forage in meadows, whereas IR[HVIRUDJHGLQIRUHVWZLWKKLJKYROXPHVRILQVHFWVDVZHOODVQHFURSKDJHV0DUWHQV SUHIHUUHGVPDOOHUZKLOHIR[HVSUHIHUUHGODJHULQVHFWV:HFRQFOXGHWKDWLQVHFWVIRXQG in faeces might play an important role in understanding food and habitat relationships between sympatric predators.

Introduction very similar (Fedriani et al  5HPDLQV RI LQVHFWV RU EURDGO\ VSHDNLQJ $ ODUJH QXPEHU RI FDUQLYRUH VSHFLHV FRH[LVW LQ invertebrates, frequently occur in carnivore WKHVDPHWHUULWRU\DQGH[SORLWVLPLODUUHVRXUFHV faeces. Their occurrence in the consumed food HJ IRRG KDELWDW  6FDW DQDO\VLV KHOSV XV WR depends on the predator (Kauhala et al  understand feeding behaviour and habitat prefer- Fedriani et al5XVVHOO 6WRUFK RQ ences as well as niche separation of sympatric DYDLODEOH UHVRXUFHV =LHOLQVNL et al  5H\- VSHFLHV -ĕGU]HMHZVND  -ĕGU]HMHZVNL   QROGV $HELVKHU DQGRQVHDVRQ /DQV]NL Most dietary analyses focus on the frequency .|UPHQGL*HQRYHVLet al 7KLV distribution, fresh biomass and source of prey. NLQGRIIRRGLVPRVWO\VXSSOHPHQWDU\ &DUYDOKR In case of opportunistic carnivores such as mar- *RPHV EXWPD\DOVRFRQVWLWXWHDYDOX- tens (Martes martes, M. foina  DQG WKH UHG IR[ DEOH SURSRUWLRQ RI IRRG IRU IR[HV EDGJHUV DQG (Vulpes vulpes  WKH PDLQ VRXUFHV RI IRRG DUH raccoon dogs (Kauhala et al  )HGULDQL et

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al/DQV]NL .|UPHQGL UHDFKLQJ 1DWLRQDO 3DUN 213  ƒ1 ƒ(  DQG XSWRRIIUHVKELRPDVVRIIDHFHV6WRPDFK *RUFH1DWLRQDO3DUN *13  ƒ1ƒ(  DQDO\VLVRIIR[HVVKRZHGWKDWXSWRRIWKH located in southern Poland, each differing in fresh weight of insects can be digested by this KDELWDWXVHDQGODQGFRYHU213LVDVPDOOGHFLG- FDUQLYRUH 5H\QROGV $HELVKHU  XRXVIRUHVWFRPSOH[RIDSSUR[LPDWHO\KDRI In spite of the high frequency and, some- patchily distributed woods, surrounded by many times, important role in dietary supplementation, semi-natural and anthropogenic habitats, whereas information on insect composition and group *13FRQVLVWVRIDODUJHEHHFKDQGVSUXFHIRUHVW SUHIHUHQFHVLVVFDUFH.RçHQi  UHFRJQL]HG FRPSOH[ KD ZLWKVPDOOSDWFKHVRIVHPL FDUDELGEHHWOHVLQRIVDPSOHVRIIR[VFDWV QDWXUDO PHDGRZV 5HG IR[ Vulpes vulpes  DQG The treatment of this fraction as ‘unrec- both species of martens, i.e. beech or stone RJQL]DEOH· VHHPV WR EH DQ RYHUVLPSOLÀFDWLRQ marten (Martes foina DQGSLQHPDUWHQ Martes /DQV]NL DQG .|UPHQGL   DQG 5XVVHOO martes  DUH FRPPRQO\ GLVWULEXWHG PHVRFDUQL- DQG 6WRUFK   ZKR LGHQWLÀHG VRPH LQVHFW vores in Poland. They are abundant in both study remains to species level, indicated a rich vari- areas and as it is not possible to differentiate ety of insect groups in particular samples and between the scats of marten species using visible EHWZHHQ DQDO\VHG FDUQLYRUHV 2QO\ 6NãRGRZVNL features, we recorded their scats as belonging to DQG 3RVãXV]Q\   DQDO\VHG GLHWDU\ GLIIHU- PDUWHQVSS :LHU]ERZVNDet al  ences between pine and stone martens on the 6FDWV ZHUH FROOHFWHG IURP WUDQVHFWV DORQJ basis of composition. IRUHVWU\ URDGV 7UDQVHFWV ZHUH ZDONHG WZLFH D Individual invertebrates are generally restricted month and only fresh scats were collected. To to a given area and many factors such as land- eliminate the preferences of a few carnivores, use pattern, vegetation structure or moisture can ZHUDQGRPO\VHOHFWHGVDPSOHVIURPDODUJHU separate different species groups, and are, subse- collection for a particular analysis. The total quently, of great use in bioindication (Boscaini et QXPEHURIFROOHFWHGIDHFHVZDVLQ213DQG al  5DLQR  1LHPHOl   ,QIRUPDWLRQ LQ*136FDWVZHUHFROOHFWHGGXULQJGLIIHU- RQWKHSUHVHQFHRIVSHFLÀFLQVHFWVLQWKHGLHWRI ent seasons and from different localities. Fresh carnivores might be useful for assessing not only VFDWV ZHUH SODFHG LQ SODVWLF EDJV DQG IUR]HQ DW food preferences, but also habitat preferences and ²ƒ&DQGZHUHSUHSDUHGXVLQJWKHPHWKRGRO- the behaviour of predators. On the basis of varia- RJ\ RI *RV]F]\ĸVNL   DQG 5H\QROGV DQG tion in the composition of invertebrates in the diet $HELVFKHU   of carnivores, we might be able to separate niche Faeces were rinsed with tap water through dimensions such as a meadow–forest gradient, D VLHYH ZLWK D PHVK VL]H RI ² PP 0LFUR- nocturnal–diurnal activity, etc. scopic and macroscopic fragments were sepa- Our main goal was to show variation in food rated into food categories. The dry mass of each preferences and invertebrate species composi- food category was measured. remains tion in relation to habitat use by two sympatric ZHUHWKHQVHSDUDWHGLQWRRUGHUVDQGLGHQWLÀHGWR FDUQLYRUHV IR[HV DQG SLQH DQG VWRQH PDUWHQV species level using reference insect collections One of our aims was to show that high variation from each region. The number of specimens was in invertebrate composition and abundance in counted for each species on the basis of replica- the food of carnivores depends not only on food EOHSDUWVRIWKHH[RVNHOHWRQVXFKDVWKHKHDGWKH DYDLODELOLW\ EXW DOVR RQ VSHFLÀF SUHIHUHQFHV RI pronotum, abdominal appendages and legs. the consumers in the type of supplementary food 7KH H[SHFWHG VSHFLHV QXPEHU LQ WKH HQWLUH consumed. data set was estimated using the Bootstrap esti- PDWRU RI VSHFLHV ULFKQHVV 6PLWK  YDQ %HOOH   Material and methods m Sbot = Sobsȸ ²pk The collection of carnivore faeces was car-

ULHGRXWLQ²LQWZRDUHDVLH2MFyZ where Sbot is estimated species richness, Sobs is

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Fig. 1. Expected number of insect species collected in the carnivores faeces using the Bootstrap estimator (± SD) of species richness.

total number of observed species in all samples Fig. 2. A biplot of the first two canonical axes of a cor- pooled, p is the proportion of samples that k respondence analysis of insect species (M) in relation contain species k, and m is the total number to environmental variables. Arrows represent environ- of samples. The randomisation of samples was mental dummy variables: main predators (marten and SHUIRUPHG XVLQJ WKH SURJUDPPH (VWLPDWH6  fox) and different landscapes (GNP = Gorce National &ROZHOO  Park, ONP = Ojcow National Park). Biomass of the most frequent species was calculated using the formula in-DURVLN   Results B a :HZHUHDEOHWRGHWHUPLQHVSHFLHVRIDUWKUR- where a is mean body length an individual. SRGVPDLQO\EHHWOHVDQGZDVSV 7DEOH Their The selection of of a particular car- frequencies varied between groups and samples. nivore was evaluated on the basis of the relative :H ZDQWHG WR GHWHUPLQH WKH H[SHFWHG QXPEHU abundance of beetle species in a given commu- of species in the area. The species rarefaction nity. curve, based on the bootstrap analysis, indicates To determine the relationship between car- WKDW WKH PDMRULW\ RI WKH H[SHFWHG IDXQD LQ WKH nivores and the insect community collected area was collected )LJ  from the faeces UHGXQGDQF\ DQDO\VLV 5'$  To determine the relationship between the DQG FDQRQLFDO FRUUHVSRQGHQFH DQDO\VLV &&$  arthropod species collected and large carnivore ZHUHXVHG WHU%UDDNWHU%UDDN äPLODXHU consumption, a canonical correspondence analy-  5'$UHYHDOVLIWKHUHH[LVWVDOLQHDUUHOD- VLV ZDV XVHG 7KH RUGLQDWLRQ GLDJUDP )LJ   tionship between response variables (insect indicated that the variation in species abun- FRPPXQLW\ VWUXFWXUH SDUDPHWHUV  DQG SUHGLFWRU GDQFHV DPRQJ UHSOLFDWHV IDHFHV  ZDV DVVRFL- YDULDEOHVV FDUQLYRUHV DQG VLWH  7KH XQLPRGDO ated mainly with carnivore species and habitat response model was selected in order to study FKDUDFWHULVWLFV DOWLWXGHDQGODQGXVHW\SH 7KH particular relationships between species and ÀUVWIRXUFDQRQLFDOD[HVGHVFULEHGRIWKH environmental factors, which are included in the YDULDQFHRIWKHVSHFLHVGDWDDQGRIWKHVSH- &&$GXHWRWKHIDFWWKDWPD[LPXPDEXQGDQFH FLHV²HQYLURQPHQWDOUHODWLRQ 7DEOH 7KHLQVHFW occurs under optimal environmental conditions. species distribution on the ordination diagram Insect abundances were log(n WUDQVIRUPHG FDQ EH FODVVLÀHG LQWR IRXU JURXSV LQ UHODWLRQ WR DYRLG WKH XQGXH LQÁXHQFH RI RXWOLHUV RQ WKH WR FDUQLYRUH DQG QDWLRQDO SDUN 6SHFLHV LQ WKH RUGLQDWLRQ :H XVHG WKH SURJUDPPH &$12&2 upper right of the diagram are mainly consumed YHU WHU%UDDN äPLODXHU  E\ IR[HV LQ WKH IRUHVW 0RVW RI WKHVH EHORQJ WR

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the carabid beetle genera and Pteros- IR[HV PRUH IUHTXHQWO\ HDW GHFD\LQJ PHDW 6SH- tichus, which were the most frequently collected cies in the lower right cluster are also consumed WD[DLQWKHGDWDVHW,QDGGLWLRQWKHSUHVHQFHRI E\ IR[HV EXW LQ WKH PHDGRZ KDELWDW 7KHUH DUH scavengers such as Silpha atrata indicates that two large species in this group: the nest building

Table 1. The frequency of occurrence (freq) and abundance (abund) of insects in marten and fox faeces collected in Gorce National Park (GNP) and Ojcow National Park (ONP).

Marten GNP Marten ONP Fox GNP Fox ONP

freq abund freq abund freq abund freq abund

01 0.00 0 0.17 4 0.00 0 0.13 1 02 Amara sp. 0.00 0 0.08 12 0.00 00.00 0 03 Aphodius sphacelatus 0.00 00.00 00.00 0 0.13 2 04 Araneus sp. 0.17 1 0.08 1 0.13 1 0.00 0 05 Bombus sp. 0.00 0 0.17 4 0.00 00.00 0 06 Byrrhus sp. 0.00 0 0.08 2 0.00 00.00 0 07 Carabus auronitens 0.17 2 0.00 0 0.25 2 0.25 2 08 Carabus cancelatus 0.00 00.00 00.00 0 0.13 1 09 Carabus coriaceus 0.00 00.00 0 0.13 1 0.00 0 10 Carabus glabratus 0.17 1 0.00 0 0.13 1 0.25 2 11 Carabus linnei 0.00 0 0.17 3 0.13 1 0.00 0 12 Carabus violaceus 0.00 00.00 0 0.13 1 0.13 1 13 Melasoma sp. 0.08 1 0.00 00.00 00.00 0 14 Cychrus craboides 0.00 00.00 00.00 0 0.13 1 15 Cychrus attenuatus 0.17 1 0.00 00.00 00.00 0 16 Forficula auriculata 0.00 0 0.25 39 0.00 00.00 0 17 Geotrupes stercorosus 0.17 3 0.33 5 0.25 2 0.25 18 18 Julus sp. 0.00 0 0.08 2 0.13 1 0.00 0 19 Necrophorus sp. 0.00 0 0.08 1 0.00 0 0.13 1 20 Orthoptera sp. 0.00 0 0.08 2 0.00 00.00 0 21 Otiorhynchus ovatus 0.00 0 0.08 1 0.00 00.00 0 22 Otiorhynchus sp. 0.00 00.00 00.00 0 0.13 1 23 Poecilus versicolor 0.17 1 0.08 9 0.00 00.00 0 24 Pseudoophonus rufipes 0.00 0 0.08 1 0.00 00.00 0 25 Pterostichus brumeisteri 0.33 3 0.00 00.00 0 0.13 1 26 Pterostichus melanarius 0.00 00.00 00.00 00.00 0 27 Pterostichus niger 0.00 00.00 0 0.13 1 0.13 1 28 Selatosomus aeneus 0.00 00.00 00.00 0 0.13 1 29 Silpha atrata 0.00 0 0.08 2 0.25 2 0.13 2 30 Vespa crabro 0.33 2 0.17 5 0.13 1 0.13 1

Table 2. Summary of canonical correspondence analysis (CCA) eigenvalues, and the cumulative percentage of variance explained on the first four canonical axes. Significance of canonical axes identified using the Monte Carlo permutation test.*p < 0.1.

Axis1 Axis 2 Axis 3 Axis 4

Eigenvalues 0.239 0.142 0.130 4.438 Species-environment correlations 0.892 0.836 0.686 0.818 Cumulative percentage variance of species data 6.80 12.2 15.4 18.3 of species-environment relation 32.0 57.3 72.2 86 F Significance of first canonical axis 1.98* Significance of all canonical axes 1.22*

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Fig. 3. Correlation biplots based on a redundancy Fig. 4. Mean niche breadths of martens and foxes analysis of the species groups and community structure according to the Shannon-Wiener index.*p < 0.05 parameter data from the faeces of martens and foxes in the two national parks (ONP and GNP). Names of the groups indicate insect preferences (necropha = scav- $UHGXQGDQF\DQDO\VLV )LJ UHYHDOHGWKDW engers, forest = forest species, eurytopi = eurytopous most of the variation of arthropod communities insects, carnivor = carnivores, omnivoro = omnivorous insects, phytophag = plant eaters, meadow = meadow and the abundances of some characteristic eco- preferring insects, nest = nest building insects) and ORJLFDOJURXSVZDVH[SODLQHGE\WKHSUHGDWRUVSH- community structure parameters (rich = species rich- FLHV IR[RUPDUWHQ 7KHÀUVWIRXUD[HVH[SODLQHG ness, abund = sample abundance). RIWKHFRPPXQLW\YDULDQFHZKLOH of the community indices are related to environ- mental factors such as carnivore identity and site Vespa crabro and the dung beetle Geotrupes ORFDWLRQ 7DEOH 7KHÀUVWDQGDOOIRXUFDQRQL- stercorosus. FDOD[HVDUHVLJQLÀFDQWDWp  0RQWH&DUOR 7KH XSSHU OHIW FOXVWHU FRQWDLQV VPDOOVL]HG SHUPXWDWLRQ WHVW  7KHVH GXPP\ YDULDEOHV LH consumed by marten, which are char- PDUWHQ DQG IR[ ZHUH FRUUHODWHG ZLWK WKH ÀUVW DFWHULVWLFRIWKHIRUHVWÁRRUIRUH[DPSOHWKHQHVW FDQRQLFDOD[LVZKLFKLQGLFDWHVSULPDU\VLJQLÀ- building earwig, )RUÀFXOD DXULFXODWD, which cance of these variables. The scatter plot of the always occurred in faeces in large numbers. The data indicates two groups of arthropods indi- lower left group contains mostly meadow spe- ces. Martens prefer meadow and nest-building cies such as a nest building Bombus sp. or large VSHFLHV ZKHUHDV IR[HV SUHIHU IRUHVW VSHFLHV RI orthopterans. greater biomass and necrophages. There was a

Table 3. Summary of redundancy analysis (RDA) eigenvalues, cumulative percent of variance explained on the first four canonical axes. Significance of canonical axes identified using the Monte Carlo permutation test. *p < 0.05.

Axis1 Axis 2 Axis 3 Axis 4

Eigenvalues 0.156 0.054 0.019 0.002 Species-environment correlations 0.665 0.484 0.462 0.187 Cumulative percentage variance of species data 16.3 22.0 24.0 24.2 of species-environment relation: 67.0 90.3 98.6 99.6 F Significance of first canonical axis 5.45* Significance of all canonical axes 1.80*

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Fig. 6. The distribution of biomass in relation to size classes (1: 5–15 mm, 2: 15–25 mm, 3: 25–35 mm) of marten and fox food. Significant differ- ences between pairs of size groups are marked with asterisks (*).

Fig. 5. Relationship between the abundance of ground beetles in the forest community and the abundance of carabid beetles in (A) marten and (B) fox faeces. tions (O’Mahony et al1RZDN $V they are opportunistic feeders, their food prefer- HQFHV UHÁHFW ORFDO SUH\ DYDLODELOLW\ 6LPXOWDQH- higher abundance and richness of insect species ously these predators may switch to alternative IRXQGLQWKHVFDWVFROOHFWHGLQ213 prey (Ben-David et al%DOWUŗQDLWē  7RWDOQLFKHEUHDGWKVRIPDUWHQVDQGIR[HVLQ A large number of researchers have stud- relation to arthropod consumption are presented LHG GLHWDU\ H[SORLWDWLRQ E\ V\PSDWULF VSHFLHV, LQ)LJ)R[IRRGLVPRUHGLYHUVHDFFRUGLQJWR RI ZKLFK UHG IR[HV DQG PDUWHQV DUH IUHTXHQW the 6KDQQRQ:LHQHULQGH[ (t p   H[DPSOHV 6HUDÀQL /RYDUL/DQV]NLet al. There was a strong positive correlation be- %LUNV5XVVHOO 6WRUFK:LHU]- tween the relative abundance of ground beetles ERZVND et al  6LGRURYLFK et al   in the forest community and the occurrence of ,QVSLWHRIWKHIDFWWKDWWKH\VKRZVLJQLÀFDQWO\ VSHFLHVLQWKHIDHFHVRIPDUWHQ )LJD 6XFKD overlapping habitat and food preferences, some UHODWLRQVKLSZDVQRWREVHUYHGIRUIR[HV )LJE  authors believe that these carnivores do not com- This indicates that martens are more opportunis- pete. They share the same habitat and similar tic feeders)R[HVDUHPRUHVHOHFWLYHSUHIHUULQJ food resources but demonstrate minor discrep- only certain species, which are not necessarily DQFLHVWKDWDOORZWKHPWRFRH[LVW -ĕGU]HMHZVND most abundant.  -ĕGU]HMHZVNL  /фSH]0DUWLQ   :KHQZHFRPSDUHVL]HFODVVHVLQIRRGFRQ- $OWKRXJKPDUWHQVDQGIR[HVDUHIRRGJHQHUDOLVWV VXPHG E\ IR[HV DQG PDUWHQV WKH GLVWULEXWLRQV there are distinct insect groups that appear more DUH GLIIHUHQW )LJ   )R[HV FRQVXPH VLJQLÀ- frequently in their individual diets. Yet, arthro- cantly more large carabids (F p   SRGVFRQVWLWXWHDVLJQLÀFDQWSDUWRIIRRGRIERWK ZKLOH WKH GLHW RI PDUWHQV LQFOXGHV VLJQLÀFDQWO\ species (Molsher et al&DUYDOKR *RPHV more smaller beetles (F p  .  0RVWRIWKHUHFHQWVWXGLHVDQDO\VLQJWKH IRRGRIPDUWHQVDQGIR[HVWUHDWLQYHUWHEUDWHVDV an unrecognisable fraction and focus only on its Discussion proportion in the whole diet. Few authors have observed that invertebrates might constitute a )R[HVDQGPDUWHQVDUHJHQHUDOLVWSUHGDWRUVZLWK ODUJH SURSRUWLRQ RI WKH IRRG RI IR[HV DQG PDU- broad food ranges and wide geographic distribu- WHQV -ĕGU]HMHZVNL et al  /DQV]NL et al.

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 7KHFRQWULEXWLRQRILQYHUWHEUDWHVGHSHQGV YHU\ VLJQLÀFDQW LQ UHODWLRQ WR DOO WKH IRRG FRQ- mainly on season and location. Insects constitute VXPHG 5H\QROGV  $HELVKHU   6XFK D an important dietary supplement of carnivores food type in WKHGLHWRIIR[HVZDVDOVRFRQÀUPHG during spring and summer in temperate areas E\ RWKHU DXWKRUV /OR\G  .RçHQi   /DQV]NLet al:LHU]ERZVNDet al  However, both authors believe that although the or during the whole year in warmer areas such beetles appear more frequently in the diet of the DV WKH 0HGLWHUUDQHDQ 6HUDÀQL  /RYDUL  UHGIR[WKHLUFRQWULEXWLRQLVRIOLWWOHLPSRUWDQFH *HQRYHVLet al  ZLWKVRPHVHDVRQDOH[HSWLRQV. &&$DQDO\VLVUHYHDOHGWKDWPRVWRIWKHYDULD- +RZGRZHH[SODLQ the presence of many car- tion in invertebrate species in faeces is described rion-feeding insects in WKHIDHFHVRIIR[HV")R[HV E\WKHIDFWRU¶VSHFLHV· LHIR[DQGPDUWHQ 7KLV presumably feed on carrion more frequently than information is very useful when comparing niche PDUWHQVDQGWKHSUHVHQFHRIVXFKLQVHFWVLQIR[ RYHUODSDQGFRPSHWLWLYHH[FOXVLRQRIWKHFDUQL- faeces indicates the scavenging behaviour of vores. Preferences of particular insect species this predator. However, some authors have also may indicate food selectivity, but in most cases demonstrated that martens feed on carcasses it is a compromise between competitive effect (Ben-David et al6LGRURYLFKet al and food availability. Only few insect species %DOWUŗQDLWē  

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