Diet selection by hares (Lepus europaeus) in arable land and its implications for habitat management

Abstract

Populations of European hares (Lepus europaeus) have experienced a dramatic decline throughout Europe in recent decades. European hares are assumed to prefer weeds over arable crops, and weed abundance was reduced by the intensification of agriculture. Therefore, modern agriculture has been blamed as a major factor affecting European hare populations. However, it is questionable whether European hares select weeds at all, as previous studies had major methodological limitations. By comparing availability and use of plants with Chesson’s Electivity Index, we investigated whether the European hare actually feeds selectively on different plants in arable land. Food availability and use were dominated by cultivated crops (e.g. winter wheat, spring barley and sugar beet). Diet selection analysis revealed that in autumn and winter, European hares predominantly preferred cultivated crops (winter wheat) and food items provided by hunters (tubers of sugar beet and carrot). In spring and summer, apart from soy, only weeds (e.g. clover and corn poppy) were positively selected, especially after cereal crops were harvested. We suggest that the decline in European hare populations throughout Europe was facilitated by the decrease in weed abundance. Wildlife-friendly set-asides in arable land have the potential to reconcile the European Union’s Common Agricultural Policy with wildlife conservation.

Appendices

Appendix 1

Plant taxa available to the sampled hares. The availability reflects the percentage of the sum of plant cover percentages for each individual taxon according to season in the MPFA of the 117 sampled hares; + indicates that this taxon was present, yet <0.1%.

Period	Scientific name	Percent
February	Sinapis arvensis	28.03
	Sinapis arvensis/Phacelia tanacetifolia	14.55
	Calamagrostis epigejos	11.51
	Triticum aestivum	5.38
	Fabaceae	4.22
	Avenella flexuosa	4.22
	Pisum sativum/Phacelia tanacetifolia
	Sinapis arvensis	3.73
	Pisum sativum	3.22
	Asparagus officinalis	2.88
	Arrhenatherum elatius	2.54
	Hordeum vulgare	1.91
	Medicago sativa	1.87
	Phragmites australis	1.70
	Stellaria media	1.60
	Bromus sterilis	1.48
	Zea mays	1.17
	Arrhenatherum elatius/Calamagrostis epigejos	1.08
	Poa pratensis	1.04
	Bromus japonicus	<1
	Bromus sp.	<1
	Cirsium arvense	<1
	Dactylis glomerata	<1
	Erigeron annuus	<1
	Fragaria vesca	<1
	Gramina sp.	<1
	Helictotrichon pubescens	<1
	Juncus filiformis	<1
	Lolium perenne	<1
	Matricaria recutita	<1
	Muscus sp.	<1
	Pisum sativum/Sinapis arvensis	<1
	Panicum miliaceum	<1
	Secale cereale	<1
	Seedlings	<1
	Solidago canadensis	<1
	Solidago virgaurea	<1
	Sorghum sudanense	<1
	Agropyron repens	+
	Allium cepa	+
	Avena sativa	+
	Beta vulgaris	+
	Brassica oleracea	+
	Bromus ramosus	+
	Calamagrostis sp.	+
	Carex acutiformis	+
	Carex otrubae	+
	Carex sp.	+
	Composita	+
	Cornus sanguinea	+
	Cynodon dactylon	+
	Daucus carota	+
	Galium verum	+
	Holcus mollis	+
	Juncus sp.	+
	Lamium purpureum	+
	Lythrum salicaria	+
	Malus domestica	+
	Mentha sp.	+
	Papaver rhoeas	+
	Poaceae sp.	+
	Rumex obtusifolius	+
	Syringa vulgaris	+
	Urtica dioica	+
May	Triticum aestivum	52.39
	Hordeum vulgare	21.84
	Beta vulgaris	5.85
	Pisum sativum	1.95
	Secale cereale	1.92
	Medicago sativa	1.63
	Helianthus annuus	1.61
	Calamagrostis epigejos	1.44
	Zea mays	1.24
	Phaseolus vulgaris	1.03
	Agropyron repens	<1
	Allium cepa	<1
	Arrhenatherum elatius	<1
	Asparagus officinalis	<1
	Avena fatua	<1
	Avena sativa	<1
	Avenella flexuosa	<1
	Bromus japonicus	<1
	Bromus sterilis	<1
	Cirsium arvense	1
	Dactylis glomerata	<1
	Daucus carota	<1
	Fabaceae	<1
	Fragaria vesca	<1
	Gramina sp.	<1
	Lathyrus sativus	<1
	Lolium perenne	<1
	Phacelia tanacetifolia	<1
	Phragmites australis	<1
	Poa pratensis	<1
	Sinapis arvensis	<1
	Solidago canadensis	<1
	Solidago virgaurea	<1
	Urtica dioica	<1
	Achillea millefolium	+
	Agrostis spica-venti	+
	Amaranthus retroflexus	+
	Anthriscus sylvestris	+
	Arctium lappa	+
	Arctium tomentosum	+
	Artemisia vulgaris	+
	Atriplex patula	+
	Brachypodium pinnatum	+
	Bromus hordeaceus	+
	Bromus ramosus	+
	Capsella bursa-pastoris	+
	Cardaria draba	+
	Carduus acanthoides	+
	Carex acutiformis	+
	Chenopodium alba	+
	Conium maculatum	+
	Consolida regalis	+
	Convolvulus arvensis	+
	Conyza canadensis	+
	Cornus sanguinea	+
	Cynodon dactylon	+
	Descurainia sophia	+
	Dipsacus fullonum	+
	Epilobium roseum	+
	Euonymus europaea	+
	Festuca ovina	+
	Festuca rubra	+
	Galium aparine	+
	Galium mollugo	+
Glycine max	+
Hordeum murinum	+
Hypochaeris radicata	+
Lamium amplexicaule	+
Leontodon autumnalis	+
Linaria vulgaris	+
Lolium sp.	+
Lotus corniculatus	+
Medicago sativa/Trifolium incarnatum	+
Matricaria recutita	+
Medicago lupulina	+
Orchis sp.	+
Panicum miliaceum	+
Papaver rhoeas	+
Poa annua	+
Poaceae and Fabaceae	+
Polygonum aviculare	+
Rubus caesius	+
Rumex obtusifolius	+
Salvia sylvestris	+
Shrubs	+
Silene vulgaris	+
Solanum dulcamara	+
Solanum tuberosum	+
Sonchus oleraceus	+
Sonchus sp.	+
Spinacia oleracea	+
Stellaria media	+
Symphytum officinale	+
Syringa vulgaris	+
Taraxacum officinale	+
Trifolium alexandrinum	+
Trifolium pratense	+
Trifolium repens	+
Trifolium sp.	+
Trifolium suaveolens	+
Tripleurospermum inodorum	+
Veronica persica	+
Vicia faba	+
Vicia sativa	+
Viola arvensis	+
August	Beta vulgaris	27.27
	Zea mays	13.30
	Helianthus annuus	9.23
	Medicago sativa	8.24
	Solanum tuberosum	6.34
	Asparagus officinalis	4.37
	Glycine max	3.40
	Avenella flexuosa	2.70
	Turf	2.16
	Calamagrostis epigejos	1.98
	Phaseolus vulgaris	1.92
	Cirsium arvensee	1.84
	Chenopodium alba	1.73
	Hordeum vulgare	1.35
	Triticum aestivum	1.22
	Daucus carota	1.20
	Fabaceae	1.12
	Pisum sativum	1.01
	Agropyron repens	<1
	Agrostis spica-venti	<1
	Allium cepa	<1
	Amaranthus retroflexus	<1
	Arrhenatherum elatius	<1
	Bromus sterilis	<1
	Convolvulus arvensis	<1
	Conyza Canadensis	<1
	Dactylis glomerata	<1
	Dipsacus fullonum	<1
	Erigeron annuus	<1
	Fragaria vesca	<1
	Lactuca serriola	<1
	Lolium perenne	<1
	Lolium sp.	<1
	Phacelia tanacetifolia	<1
	Phragmites australis	<1
	Poa pratensis	<1
	Solidago canadensis	<1
	Solidago virgaurea	<1
	Solidago virgaurea	<1
	Trifolium repens	<1
	Trifolium saxatile	<1
	Urtica dioica	<1
	Acer negundo	+
	Achillea millefolium	+
	Amaranthus alba	+
	Arctium lappa	+
	Artemisia vulgaris	+
	Brachypodium pinnatum	+
	Bromus japonicus	+
	Bromus ramosus	+
	Bromus sp.	+
	Calystegia sepium	+
	Carduus acanthoides	+
	Carduus sp.	+
	Carex acutiformis	+
	Carex otrubae	+
	Carex pilulifera	+
	Carlina vulgaris	+
	Conium maculatum	+
	Cornus sanguinea	+
	Datura stramonium	+
	Deschampsia caespitosa	+
	Descurainia sophia	+
	Echium vulgare	+
Epilobium roseum	+
Erigeron sp.	+
Euonymus europaea	+
Fagopyrum esculentum	+
Fallopia convovulus	+
Festuca rubra	+
Fraxinus excelsior	+
Galium aparine	+
Galium mollugo	+
Galium verum	+
Helictotrichon pubescens	+
Hypochaeris radicata	+
Inula conyza	+
Juncus filiformis	+
Lathyrus sativus	+
Linaria vulgaris	+
Lotus corniculatus	+
Matricaria recutita	+
Mentha aquatica	+
Panicum miliaceum	+
Papaver rhoeas	+
Pastinaca sativa	+
Phleum pratense	+
Poaceae sp.	+
Polygonum aviculare	+
Rubus caesius	+
Rumex obtusifolius	+
Secale cereale	+
Securigera varia	+
Senecio vulgaris	+
Setaria verticillata	+
Setaria viridis	+
Solanum dulcamara	+
Sonchus oleraceus	+
Spinacia oleracea	+
Symphytum officinale	+
Trifolium incarnatus/suaveolens	+
Taraxacum officinale	+
Trifolium alexandrinum	+
Trifolium suaveolens	+
Tripleurospermum inodorum	+
Tussilago farfara	+
Vicia faba	+
Vicia sativa	+
November	Phacelia tanacetifolia	20.43
	Sinapis arvensis	17.60
	Triticum aestivum	13.23
	Avenella flexuosa	6.96
	Calamagrostis epigejos	6.87
	Medicago sativa	5.79
	Cirsium arvense	3.16
	Bromus sterilis	2.96
	Trifolium pretense	2.78
	Lathyrus sativus	2.74
	Arrhenatherum elatius	1.96
	Vicia sativa	1.70
	Solidago virgaurea	1.53
	Pisum sativum	1.26
	Conyza canadensis	1.19
	Lolium perenne	1.02
	Seedlings	1.02
	Acer negundo	<1
	Agropyron repens	<1
	Agrostis spica-venti	<1
	Amaranthus retroflexus	<1
	Artemisia vulgaris	<1
	Atriplex patula	<1
	Avena fatua	<1
	Bromus japonicus	<1
	Bromus sp.	<1
	Dactylis glomerata	<1
	Dipsacus fullonum	<1
	Festuca rubra	<1
	Leontodon autumnalis	<1
	Lolium sp.	<1
	Lotus corniculatus	<1
	Muscus sp.	<1
	Phragmites australis	<1
	Poa pratensis	<1
	Rubus caesius	<1
	Sambucus nigra	<1
	Trifolium repens	<1
	Tripleurospermum inodorum	<1
	Zea mays	<1
	Achillea millefolium	+
	Arctium lappa	+
	Arctium tomentosum	+
	Beta vulgaris	+
	Brachypodium pinnatum	+
	Bromus ramosus	+
	Capsella bursa-pastoris	+

Appendix 2

List of plant taxa found in the hares’ stomachs (N=117). Sample periods being February (N=18 plants), May (N=25 plants), August (N=24 plants) and November (N=20 plants). Percentages shown as proportions of the sum of each individual plant taxon per period; + indicates that this taxon was present, yet <0.1%.

Season	Scientific name	Percent
February	Triticum aestivum	57.07
	Beta vulgaris	24.14
	Medicago sativa	9.87
	Sinapis arvensis	2.78
	Malus domestica	2.04
	Daucus carota	1.85
	Stellaria media	1.44
	Zea mays	0.32
	Galium verum	0.14
	Lamium purpureum	0.14
	Juncus capitatus	0.12
	Avenella flexuosa	+
	Bark	+
	Cirsium sp.	+
	Cornus sanguinea	+
	Dactylis glomerata	+
	Juncus sp.	+
	Taraxacum officinale	+
May	Triticum aestivum	50.12
	Hordeum vulgare	12.93
	Glycine max	11.32
	Trifolium pratense	7.71
	Beta vulgaris	5.44
	Papaver rhoeas	3.42
	Trifolium repens	2.52
	Secale cereale	1.83
	Taraxacum officinale	1.7
	Medicago sativa	0.95
	Dactylis glomerata	0.78
	Pisum sativum	0.58
	Rubus caesius	0.25
	Arrhenatherum elatius	0.13
	Agropyron repens	+
	Bromus hordeaceus	+
	Bromus japonicus	+
	Bromus sterilis	+
	Capsella bursa-pastoris	+
	Cirsium arvense	+
	Convolvulus arvensis	+
	Poa annua	+
	Polygonum aviculare	+
	Stellaria media	+
	Trifolium pretense	+
	Trifolium repens	+
	Zea mays	+
August	Medicago sativa	38.79
	Hordeum vulgare	25.49
	Beta vulgaris	13.87
	Artemisia vulgaris	4.34
	Daucus carota	2.98
	Triticum aestivum	2.45
	Taraxacum officinale	2.31
	Zea mays	2.3
	Avenella flexuosa	1.77
	Glycine max	1.44
	Agropyron repens	1.18
	Amaranthus retroflexus	1.03
	Panicum miliaceum	0.88
	Dactylis glomerata	0.64
	Polygonum aviculare	0.12
	Fagopyrum esculentum	0.1
	Lathyrus sativus	0.1
	Arrhenatherum elatius	+
	Convolvulus arvensis	+
	Hordeum murinum	+
	Lotus corniculatus	+
	Papaver rhoeas	+
	Taraxacum officinale	+
	Thymus serpyllum	+
	Trifolium incarnatus/suaveolens	+
November	Triticum aestivum	51.86
	Beta vulgaris	21.72
	Medicago sativa	17.37
	Sinapis arvensis	3.65
	Arrhenatherum elatius	1.65
	Avenella flexuosa	1.6
	Trifolium pratense	1.19
	Bromus sp.	0.52
	Dactylis glomerata	0.35
	Agropyron repens	+
	Amaranthus retroflexus	+
	Artemisia vulgaris	+
	Capsella bursa-pastoris	+
	Crataegus sp.	+
	Helictotrichon pubescens	+
	Malus domestica	+
	Phacelia tanacetifolia	+
	Plantago lanceolata	+
	Stellaria media	+
	Zea mays	+