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Mammal Bible
Report to: Mr. M. Clook Ecotoxicology Section Pesticides Safety Directorate Mallard House, Kings Pool 3 Peasholme Green York YO1 2PX CSL Contract Manager: Dr. A.D.M. Hart Central Science Laboratory Sand Hutton York YO41 1LZ Date of issue of report: November 1998
1998 Update CONTRACT PN0910/PN0919
MILESTONE REPORT Mammals and farming:
information for risk assessment J.E. Gurney, J. Perrett
D.R. Crocker & J.A. Pascual
CSL Project No. M37
2
Contents
INTRODUCTION 3
GENERAL ECOLOGICAL INFORMATION (30 SPECIES IN ALPHABETICAL ORDER) 6
DETAILED DIETARY INFORMATION (30 SPECIES IN ALPHABETICAL ORDER) 51
BIBLIOGRAPHY 207
APPENDIX: EXAMPLES OF RESULTS GENERATED BY ACCESS QUERIES 219
3
Introduction
This compendium is intended as a companion volume to the earlier volume “Birds and farming” (Buxton and Crocker, 1996). It has two broad purposes. Firstly it is intended to suggest which mammal species are most likely to be exposed to particular plant protection products. Thus information has been collected on geographical distribution of mammals on farmland, their typical feeding and breeding habitats, known associations with crops, the times of year when young are being reared, and on population status and trends. Secondly, it is intended to help calculate the likely degree to which a given species may be exposed to a given plant protection product. Using data on pesticide residues found on different food types, the likely exposure of an individual may be calculated -- either directly from data in the review giving actual daily weights of foods eaten; or indirectly by using an species’ body mass to estimate total daily food requirement and then using data on the proportion of the diet made up by a given food type. The data collected in the course of the review have been stored on a Microsoft Access database. This greatly increases their usefulness by allowing questions to be asked of the database. Thus it is a relatively simple matter to provide lists of species associated with particular habitat, crops, or foods. It may be useful to identify early or late-breeding mammals, small mammals, mammals in decline and so on. More complex queries may also be posed such as: which mammals weighing less than 500g, known to be declining in numbers, are also known to have a preference for a particular farming habitat? Some examples of the tables that may be produced using Access queries are included in the Appendix. In this edition of the compendium we have drawn on additional material collected in project PN09019 “Improving estimates of the exposure of non-target wildlife to pesticides in arable crops- a review of existing data.” Reflecting the dual function, the compendium is in two parts. 1 General Ecological Information Details of each of 30 species common on British farmland or otherwise of interest to PSD are summarised in alphabetical order. With a single page devoted to each species, the descriptions of habitat, food preferences etc. are necessarily brief. Information was included if it was relevant to risk assessment rather than any intrinsic zoological interest. The following standard format was used. Data on body weights, feeding habits and habitat were mostly taken from “The Handbook of British Mammals” (Corbet and Harris, 1991) and “A review of British Mammals” (Harris, Morris, Wray and Yalden, 1995). Estimates of population size and trends were also taken from Harris et. al. (1995). Data on status and distribution were found in “Atlas of Mammals in Britain” (Arnold, 1993).
4
[Common name] [Species] [Order] [Family]
Body Weight (g) male: : [mean (range)] female: [mean (range)] (Where possible mean weights, together with known ranges are included for both male and female members.) Young born from: [month] to [month] Geography: [Description] Recorded in: [ ] 10 Km squares in Britain [ ] % Status: [Description] Density: [Description] Population: England Scotland Wales Population trend: [Declining/Slight decline/Stable/Slight increase/Increasing] Feeding Habitat: [Description] Associated crops: [Description] Foraging Behaviour: [Description] Further notes on mammals’ use of arable land: [Description] (Data taken from a review (PN0919) of mammals and arable farming conducted by CSL for PSD) Note: Where a topic appears to be missing for a particular species, it is because no information on that topic could be found for that species. 2 Detailed Dietary Information For each of the 30 representative species we conducted a more detailed review of their typical diets. For each reference cited we attempted to describe how diet varied at different times of year, what foods were taken and in what amounts. Where possible we estimated how many grams of a particular food type might be eaten in a day and the typical energy value of the daily diet. More commonly research papers gave information on the proportion of the daily diet made up by a particular food type. Different authors calculate this proportion in different ways
5
and this is indicated in the comments section. For example, estimates of consumption may have been made from stomach contents or from faeces; some authors simply count the frequency with which certain items occur, others attempt to estimate the relative volume contributed by the different foods. The comments section also indicates if the data are from captive animals, the country where the study was carried out and other relevant information.
6
General Ecological Information (30 species in alphabetical order)
Carnivora. Mustelidae
Badger Meles meles
Young born from: Jan to: Jul
Geography: Very widespread in Britain (especially in the South) andIreland but notably absent from high ground, largeconurbations, intensively farmed areas and lowlands liableto flooding.
Feeding habitat: Feed and dig setts in a wide range of habitats from openfields, hedgerows and woodland to urban situations.Prefer mixed and plentiful food supply. Permanent pasturepreferred to arable habitat
Associated crops: Pasture, cereals, wheat, oats, barley, maize, horticulture.
Recorded in: 1800 10 km squares in Britain
Population: Increasing
Male: 11600
Female: 10100 ( 6600 -
( 9100 - 16700 )Body weight (g)
Foragingbehaviour:
Activity mainly crepuscular and nocturnal. Badgers live insocial groups of about 6 and have territory sizes of 30 ha.to 150 ha. depending upon habitat quality. Movement slowwhen foraging, pausing frequently to listen. Senses ofsmell and hearing very acute in locating food. Will dig forworms and beetles.
( 63
Outline diet: Opportunistic omnivores but earthworms are the singlemost important food. All foods taken subject to availabilitywhich include larger insects, small (young) mammals,carrion (especially in winter), amphibians, cereals andfruit. Cereals eaten mostly in late summer when dryweather makes earthworms scarce. Wheat and oatspreferred to barley
Status: Widespread, common
%)
Density: Density from <1-2 per sq km in less favourable habitatsto 20 adults per sq km in better habitats. Numbers haveincreased 76% between 1988-97.
13900 )
England:
195,000
Scotland:
25,000
Wales:
35,000
Trend:
7
Carnivora. Mustelidae
Badger Meles meles
Further notes on use made of arable land:
In a survey of badger damage carried out by questionnaire, 31% of the 2000farmers who responded reported badger damage to cereal crops (Moore et al,1998). Oats received the most damage followed by maize, wheat, barley andoilseed rape. As part of the same study, seven animals were radio-tracked inSeptember and October. Before harvesting, 39% of fixes were located in foragemaize fields, whereas after harvesting the figure had dropped to 7%. During thesame period use of pasture increased from 33% to 47%.
Badgers preferred to forage in pasture with short grass, and higher densities ofearthworms. Arable land held significantly fewer earthworms than pasture andwas increasingly used during dry periods. (Kruuk, 1978a,b, Kruuk et al, 1979,Hofer, 1988).
After large scale surveys in 1988 and 1997, Wilson et al (1997) reported thatmain setts were more likely to be found in broad-leaved woodland > hedgerows> grassland, coniferous woodland , scrub, mixed woodland > bracken and arable.
In a study of 11 radio-collared individuals on intensively farmed Sussexdownland Roper et al (1995) concluded that given the choice badgers preferredwheat preferred to barley. Wheat was taken from early May (milk stage) toNovember. Radio-collared individuals spent more than 30% of foraging time inwheat fields. Barley was taken between July and September, and only by someindividuals. Some individuals may consume as much as 1100ml grain in a nightalthough 50-250ml is more common.
Observations on 25 radio-collared individuals inhabiting a landscape comprising65% arable land, 28% pasture, and 7% scrub, spent 42% of their time foragingin pasture, 28% in scrub, 24% among wheat fields and 6% in other arable fields.There was a seasonal shift from pasture and scrub in winter and spring wheatand other arable crops in summer and autumn (Sheperdson et al, 1990).
In a study of badger damage to oats in SW England, oats became attractive atthe early milk stage (growth stage 73) beginning in late July. Damage increaseduntil the crop was harvested in mid-August reaching a maximum of 10% of thecrop flattened on one particular field.
8
Rodentia Muridae
Bank vole Clethrionomys glareolus
Young born from: Apr to: Oct
Geography: Throughout mainland Britain, and on most islands.
Feeding habitat: Very varied. Preferred habitat is mature deciduouswoodland with a thick scrub or field layer. Also foundamongst thick grass, in young tree plantations andhedgerows.
Associated crops: Grassland, forestry plantations.
Recorded in: 1064 10 km squares in Britain
Population:
Male: 26.1
Female: 21.9
Body weight (g)
Foragingbehaviour:
Active throughout day and night in summer with peaks ofactivity around dawn and dusk; less nocturnal in winter.They make runways above ground under low cover. Walkand run when foraging, using senses of smell and hearing.Only eat flesh of rosehips, discarding the skin.
( 37
Outline diet: Almost wholly herbivorous. Leaves of woody plants arepreferred to those of herbs. Fleshy fruits and seeds withsoft testa eaten when available. Other food items includefungi, moss, roots, flowers, grass, insects and worms.May strip bark and take dead leaves in winter.
Status: Abundant
%)
Density: Densities can vary tremendously due to winter breedingfrom 5 to greater than 130 per ha. In arable areas, adensity of 60 per ha. typical for hedges of reasonablequality.
England:
17,750,000
Scotland:
3,500,000
Wales:
1,750,000
9
Rodentia Muridae
Bank vole Clethrionomys glareolus
Further notes on use made of arable land:
Bank voles were wholly absent from cropped fields in southern Sweden, onlyever being found in non-crop habitat islands.(Loman, 1991).
In a study of voles in a 2 ha wood surrounded by spring wheat fields nearOxford, individuals were 3 times more likely to be found in the wood than in thesurrounding fields and very rarely found more than 20 metres into the crop(Kikkawa, 1964). A more recent similar study also confirms that few voles werecaught out of woodland habitat and none more than 80m into the crop(Fitzgibbon, 1997, see also Yloner et al 1991). The reluctance of bank voles tostray far from hedgerows in arable environments is also confirmed in trappingstudies by Pollard & Relton, 1970, Eldridge, 1971, Boone & Tinklin, 1988.However, Tew (1994) reports that as the crop cover increased through thesummer, there were frequent bank vole captures in wheat and barley fields inOxfordshire, though usually within 25m of a hedge. Plesner-Jensen (1993) alsofound that the presence of a nearby hedge significantly increased the chancesof catching bank voles in the field margins. More bank voles were caught in 2mwide conservation margins where vegetation was allowed to regeneratenaturally, than in wider 8m margins sown as grass leys.
MAFF's Boxworth project also recorded no more than one or two bank volecaptures in arable fields apart from the summer of 1984 when several fieldsrecorded between 5 and 11 captures and one field caught 26. This was a highlyunusual year and no voles were recorded in the fields in autumn after theharvest.
Bank voles prefer established hedges where there is a dense understorey(Woods et al, 1996, Boone & Tinkle, 1988).
Male home range in woodland is typically 1000-2000 sq metres, with femalesoccupying 30-80% of that area (Wolton and Flowerdew, 1985).
In a study of the value of set-aside to small mammals, Tattersall et al (1997) didnot succeed in catching any bank voles on one-year rotational set-aside. Rogers(1993) also found that set-aside was avoided by voles.
10
Lagomorpha Leporidae
Brown hare Lepus europaeus
Young born from: Feb to: Oct
Geography: Widespread throughout farmland landscapes of Britainexcept in upland areas of Scotland, N. Wales andDerbyshire where replaced by Mountain hare. Smallisolated populations of Brown hare in Ireland.
Feeding habitat: Prefer arable land where cereals predominate but withavailable grass fields for summer feeding. Shorter cropspreferred with more open vegetation. Also require woodedareas and hedgerows for resting areas during day.
Associated crops: Cereals, rape, barley, turnip, grassland.
Recorded in: 1603 10 km squares in Britain
Population: Declining
Male: 3230
Female: 3430 ( 3100 -
( 2900 - 3500 )Body weight (g)
Foragingbehaviour:
Nocturnal, but crepuscular during summer months. Homeranges possibly 20 - 40 ha. in size but do not show strongterritorial activity. Movements of up to 1.7 km recorded perday.
( 56
Outline diet: Grasses, herbs and arable crops, especially cereals inearly growth stages. Prefer wild grasses and herbs, whenavailable, to cultivated forms. In summer, herbs form bulkof diet. Grasses predominate in winter when shrubs mayalso be browsed.
Status: Common but declining
%)
Density: Very difficult to assess. Possibly up to 130 - 150 haresper square km (1.3 - 1.5 per ha.), but could be over-estimate.
3750 )
England:
572,000
Scotland:
187,250
Wales:
58,000
Trend:
11
Lagomorpha Leporidae
Brown hare Lepus europaeus
Further notes on use made of arable land:
A study of 21 radio-collared hares on French arable land recorded an averagerange size of about 100ha both in winter and summer. Cereal fields wereavoided before harvest but became more attractive as stubble (Reitz & Leonard,1994). A similar study by Marboutin & Aebischer (1996) calculated mean homerange of 20 radio-collared hares to be 190 ha. Hares made greater use ofcultivated areas than would be expected by chance and less use of non-cultivated areas (woodland, hedges, grass fields, set-aside).
In a study of hares on mixed arable farm in Hampshire, Tapper & Barnes (1986)noted that the study population preferred to feed on short crops and theirpreference for cereals declined as crops developed beyond the tillering stage(see also Pepin, 1985). Pastures were important feeding areas throughout theyear. Hares shifted their activities between fields according to crop development,in particular, moving from winter cereals in the spring to grassland at othertimes. Hares generally preferred grass without livestock (see also Barnes et al1983).
In Poland, Lewandowski & Nowakowski (1993) observed that hares tended touse different crops in proportion to their availability, apart from oilseed rapewhich they avoided and stubble fields which they preferred.
12
Rodentia Gliridae
Common dormouse Muscardinus avellanarius
18
Young born from: May to: Sep
Geography: Widespread but patchily distributed in suitable habitatsfrom mid-Wales, Leicestershire and Suffolk southwards.Scarce further north with few isolated populations.
Feeding habitat: Deciduous ancient woodland with plenty of secondarygrowth and scrub of area greater than 20 ha. Also insecondary woodland and unmanaged hedgerows. Seedsfrom hazel, sweet chestnut and beech especiallyimportant. Physical structure of woodland very importantfor arboreal pathways.
Recorded in: 352 10 km squares in Britain
Population: Declining
( 15 - 26Body weight (g)
Foragingbehaviour:
Strictly nocturnal, activity decreasing in autumn andhibernating from Oct.- Apr. Agile climbers, spendingmajority of time above ground in tree canopy and shrublayer. Travel approximately 250 m per night.
( 12
Outline diet: Varies according to season. Flowers and pollen taken inearly summer, more fruits, berries and nuts later in theyear. Also take insects. Chestnuts, acorns and hazelnutsare important prior to hibernation. In captive studies, grainis not taken but water is essential.
Status: Localised and declining
%)
Density: Occur at lower density and have lower breeding potentialthan other common rodents, typically 5 per ha ( 8 - 10 perha. in good habitat).
)
England:
465,000
Scotland:
0
Wales:
35,000
Trend:
13
Insectivora Soricidae
Common shrew Sorex araneus
8.1
Young born from: May to: Sep
Geography: Throughout mainland Britain. Not in Ireland or on manynotable islands including Isles of Scilly, Orkney, OuterHebrides, Shetland and some Inner Hebrides.
Feeding habitat: Found wherever low vegetation provides cover,particularly in thick grass and scrub, hedgerows anddeciduous woodland.
Associated crops: Permanent pasture, set-aside, agricultural headlands
Recorded in: 1429 10 km squares in Britain
Population:
( 5 - 14Body weight (g)
Foragingbehaviour:
Make runways amongst ground vegetation. Home rangesvary according to season/habitat; often in region 370 - 630sq m, up to 1100 sq m. Active day and night with approx.10 periods of continuous activity alternating with shorterperiods of rest. Peaks of activity 1000 hrs and 2200 hrs.Locate prey by probing with snout and digging down to 12cm under surface. Very acute hearing and smell. Attackhead of live prey first to immobilise and eat from headdown. Wings, legs and other unpalatable parts (slime fromslugs) discarded. Millipedes may be avoided.
( 50
Outline diet: Opportunistic predator feeding on a wide range ofcommon invertebrates, particularly earthworms, woodlice,spiders, slugs, snails and insect larvae. Although showingpreferences between prey types, items tend to be takenin proportion to their availability. Small amounts of plantmaterial including seeds are also taken.
Status: Abundant
%)
Density: Variable. 42 - 69 per ha. in deciduous woodland andgrassland at summer peak. Lower in winter, 5 - 27 per ha.
)
England:
26,000,000
Scotland:
11,500,000
Wales:
4,200,000
14
Insectivora Soricidae
Common shrew Sorex araneus
Further notes on use made of arable land:
Tew et al (1994a,b), found that both common and pygmy shrews were caughtentirely in the hedgerows during the winter months, but as crop cover increasedover the summer they were occasionally caught in the cereal field itself, thoughrarely more than 20m from a hedgerow.
In the Boxworth project, Johnson et al (1992), regularly caught shrews in arablefields but in small numbers. Forty percent of spring and summer captures werewithin 10m of a field boundary. In the first year of the project (1983) the majority(48%) of shrews were caught in hedgerows. However, as the projectprogressed, more and more individuals were trapped in open fields. In 1985 and1988, 70% of shrew captures were in open fields.
In an arable area of Sweden shrew captures were mostly confined to non-crophabitat (Loman 1991).
15
Perissodactyla Cervidae
Fallow deer Dama dama
Young born from: Jun to: Aug
Geography: Throughout England although absent in the North, south ofthe Wash and some of the SW. Present in N and E Wales,central Ireland and isolated populations throughoutScotland.
Feeding habitat: Prefer mature deciduous or mixed woodland withestablished understorey for shelter. Feed in forest rides orsurrounding agricultural land.
Associated crops: Cereals, grassland, woodland
Recorded in: 546 10 km squares in Britain
Population: Stable
Male: 67000
Female: 44000 ( 35000 -
( 46000 - 80000 )Body weight (g)
Foragingbehaviour:
Mainly crepuscular lying up in cover during day. Fallow arenon-territorial and have overlapping ranges of >50 ha.Although preferring to graze will browse during winter.Very acute senses of hearing and smell, but sight morelimited to detecting movement.
( 19
Outline diet: Prefer to graze than browse. Relatively unselective bulkfeeder. Take mainly grasses in summer with some herbsand broad leaf browse. Acorns, mast and other fruits arecommon autumn and winter foods as are bramble, holly,ivy, heather and coniferous browse.
Status: Locally common
%)
Density: Variable and in the majority of cases controlled by man.Typically, 18-43 per sq km. For parts of Essex, estimatedat one deer per 6 - 8 ha. Densities on agricultural landbetween 4.6-8.0 per sq km.
52000 )
England:
95,000
Scotland:
4,000
Wales:
1,000
Trend:
16
Perissodactyla Cervidae
Fallow deer Dama dama
Further notes on use made of arable land:
Fallow deer were studied at 3 mainly agricultural habitats in Hampshire. Theywere characterised by large open landscapes of arable fields and pastureinterspersed with small woodland copses. On average arable land amounted to48% of the area, grassland 23%, closed woodland 18% and open woodland 9%At all seasons deer used arable land and grassland significantly less often thanexpected by their availability. Although fallow deer extensively utilised smallwoodlands, the author argues that it is unlikely that they contained significantfood resources (Thirgood, 1995).
17
Artiodactyla Bovidae
Feral cattle Bos taurus
Young born from: Jan to: Dec
Geography: In Britain, there are only two feral cattle herds, one inChillingham and another on Swona, Orkney.
Feeding habitat: The two feral, free-ranging populations live in differentconditions; Chillingham Park comprises medieval pastureand ancient woods; Swona comprises maritime heath andabandoned arable and pasture land.
Associated crops: Pasture, woodland
Population: Slight increase
Male:
Female: ( -
( - 300000 )Body weight (g)
Foragingbehaviour:
Diurnal. Unlike sheep, cattle tend to maintain rate of intakeat expense of diet and digestibility.
Outline diet: Grazers. Show seasonal differences in diet but littleinformation on factors affecting choice of sward.
Density: 55 for Chillingham herd
280000 )
England:
45
Scotland:
10
Wales: Trend:
18
Artiodactyla Bovidae
Feral goat Capra hircus
Young born from: Jan to: Apr
Geography: Discrete, isolated populations on mainly mountainousareas of Scotland, Wales and Ireland. Present on manyScottish islands and 2 populations on Somerset coast.
Feeding habitat: Mostly steep mountainous habitat with cliff refuges anddwarf shrub communities for food. Prefer dry, well drainedland and can be present in woodland.
Associated crops: Rough grazing and marginal land.
Recorded in: 116 10 km squares in Britain
Population: Stable
Male: 52400
Female: 41100 ( 29000 -
( 39000 - 65000 )Body weight (g)
Foragingbehaviour:
Predominately diurnal, day spent alternatively feeding andruminating. Groups of 'billies' and 'nannies' will rangeseparately within area 0.3 - 6.5 sq km, depending onhabitat quality. Will move to lower ground during badweather. Walk and run and are extremely good climbers.Frequently browse trees on hind legs to reach under-storey. Will forage according to seasonal pattern ofavailability.
( 4
Outline diet: Will browse or graze according to local food availability.Take grasses, sedges and rushes in summer and moreshrubs, gorse and browse in winter. Will also browse andstrip bark from trees such as willow, rowan, oak and alder.
Status: Isolated populations
%)
Density: Adult densities vary 1.5 - 11.8 per sq km depending uponpopulation and time of year, typical density approx. 2 persq km.
52000 )
England:
315
Scotland:
2,650
Wales:
600
Trend:
19
Artiodactlya Bovidae
Feral sheep Ovis aries
Young born from: Mar to: May
Geography: Soay breed found on Soay and Hirta and other islandsincluding Lundy. Boreray breed confined to Boreray, St.Kilda.
Feeding habitat: Maritime heathland
Associated crops: Rough grazing and marginal land.
Population: Stable
Male:
Female: ( 13800 -
( 20100 - 25600 )Body weight (g)
Foragingbehaviour:
Diurnal; in winter most of day spent grazing; in summerspend more of day ruminating. Sheep are predominantlyor exclusively grazers. Both breeds can be selective in dietwhich shows seasonal variation. May browse during winter.
Outline diet: Almost wholly grazers. On Soay and Hirta, grasses aretaken in summer with heather increasing in the diet duringthe winter. Sheep on Boreray graze all year round.
Density: Soay; 0.9 - 2.8 sheep/ ha. Boreray approx. 12 / ha.
19800 )
England:
150
Scotland:
1,850
Wales:
100
Trend:
20
Rodentia Muridae
Field vole Microtus agrestis
Young born from: Mar to: Oct
Geography: Widespread through mainland Britain. Absent fromIreland, Channel islands, Isles of Scilly and some of theInner and Outer Hebrides.
Feeding habitat: Mainly rough, ungrazed grassland including young forestryplantations with thick grass ground cover. Low populationdensities also occur in woodlands, hedgerows, dunes andmoorland. Optimal habitat declining with increased grazingpressure from rabbits and intensive agriculture.
Associated crops: pasture, grassland, forestry.
Recorded in: 1302 10 km squares in Britain
Population: Slight decline
Male: 39.7
Female: 30.9 ( 15 -
( 15 - 42 )Body weight (g)
Foragingbehaviour:
Generally nocturnal in summer, more diurnal in winter withpeaks of activity at dawn and dusk. Short term (2 - 4hours) rhythm of activity also shown. Females haveoverlapping home ranges of approx. 200 sq m, maleslarger and more exclusive. Olfactory communication isvery important in foraging, other senses little studied.
( 46
Outline diet: Herbivorous, mainly feeding on green leaves and stemsof grasses. Some studies indicate a preference for herbsrather than grasses and sedges. Captive animals show aspring preference for more succulent grass species.
Status: Abundant
%)
Density: Very few studies due to high variability. Density of 100per ha. recorded in grassland in spring in S. England.
32 )
England:
17,500,000
Scotland:
41,000,000
Wales:
16,500,000
Trend:
21
Rodentia Muridae
Field vole Microtus agrestis
Further notes on use made of arable land:
Field voles were occasionally trapped in Aberdeenshire oilseed rape but usuallyby the edge of the field near grassy banks. Dietary analysis showed mice to beeating mainly monocotyledons (82%) and only 3% oilseed rape. (Rogers, 1990).In a two year study of small mammals on Scottish arable land and set-aside,Rogers (1993) caught 159 field voles and reports "an almost exclusivepreference for rough grassland. They were completely absent from the woodand also infrequent in set-aside and crop." Field voles prefer dense grassyhabitats in which they can conceal runways (See also Eldridge, 1971).
In a three year study of small mammals on an arable farm in Oxfordshire, Tew(1994) failed to capture any field voles away from the hedgerows around cerealfields.
Unlike bank voles, field voles showed a significant preference for newlyestablished hedgerows which had been undersown with Festuca rubra andDactylis glomerata (Woods et al, 1996)
In the Boxworth project, field voles were occasionally caught in the fields but thiswas usually restricted to areas with dense ground cover, such as patchesinfested with blackgrass (Johnson et al, 1992).
22
Carnivora Canidae
Fox Vulpes vulpes
Young born from: Feb to: Apr
Geography: Almost ubiquitous on British mainland and in Ireland.Becoming more widespread in Norfolk and Aberdeenshirebut absent from Channel Isles, Isles of Scilly and mostScottish Islands..
Feeding habitat: Very wide ranging from open agricultural land, towoodlands and urban expanses. Generally likefragmentary habitats, which offer cover and a food source.
Recorded in: 1744 10 km squares in Britain
Population: Increasing
Male: 6800
Female: 5700 ( 4200 -
( 5500 - 9300 )Body weight (g)
Foragingbehaviour:
Mainly nocturnal and crepuscular. Live in family groupswith a shared territory, often >20 ha. Will scavenge, takefruit and beetles and hunt for worms probably by hearing.Will cache food and partially bury it, returning by memorylater.
( 61
Outline diet: Omnivores. Frequently take rabbits, wood mice and fieldvoles. Bank voles and insectivores uncommon.Passerines, pigeons and game birds are the frequentlytaken bird species. Also important are a wide range ofinvertebrates (including insects and earthworms), carrionand fruit.
Status: Common and widespread
%)
Density: Typical farmland habitat of lowland England, one familygroup per sq km. Urban densities seen to vary 0.19 - 2.24family groups per sq km.
8200 )
England:
195,000
Scotland:
23,000
Wales:
22,000
Trend:
23
Carnivora Canidae
Fox Vulpes vulpes
Further notes on use made of arable land:
In a mixed farming area of S England comprising winter cereals (39%), springcereals (24%) and permanent pasture (16%), two thirds of fox diet consisted ofvertebrate prey in the range 0.3 - 3.0 kg. Small rodents, fruit and vegetableswhich are very important in other habitats amounted to less than 10% each herebecause of the presence of abundant game species; pheasant, brown hare,grey partridge and red-legged partridge (Reynolds and Tapper, 1995).
In Scotland, Hewson (1986) reported that occupied breeding dens were threetimes more numerous on agricultural land than in hill country managed forshooting red deer and twice as numerous as on heather moorland managed forred grouse. He concluded that the differences were probably due to theavailability of food.
24
Rodentia Sciuridae
Grey squirrel Sciurus carolinensis
550
Young born from: Feb to: Jul
Geography: Throughout England and Wales up to Cumbria. Alsopresent in central Scotland and central Ireland only.
Feeding habitat: Most abundant in mature broad-leafed forests of oak,beech, sweet chestnut and hazel. Also found inconiferous/broadleaf mixed woodland, hedgerows, parksand gardens and urban areas if mature trees are present.
Associated crops: Mature forestry plantations.
Recorded in: 1476 10 km squares in Britain
Population: Increasing
( 400 - 720Body weight (g)
Foragingbehaviour:
Diurnal, activity very similar to Red squirrel but lessarboreal. Individual home ranges smaller than Redsquirrels hence greater densities. Female home range 5 -6 ha. in mixed deciduous woodland, males greater. Goodhearing and sense of smell, sight suited to dim light.
( 52
Outline diet: Mainly take seed and plant buds and flowers. Prefer seed(coniferous and deciduous) but will take a wide variety ofother foods. Little evidence that eggs or chicks contributesignificantly to the diet.
Status: Common, increasing
%)
Density: Long term densities higher than for Red squirrels, usually>2 per ha., 7.4 per ha. recorded in English oak wood.
)
England:
2,000,000
Scotland:
200,000
Wales:
320,000
Trend:
25
Rodentia Muridae
Harvest mouse Micromys minutus
7
Young born from: May to: Oct
Geography: Throughout England except in North East. Records fromcoastal strip of Wales and from Scotland probablyrepresent isolated populations. Absent from Ireland.
Feeding habitat: Amongst tall grasses and monocotyledonous crops, feedon seeds and insects. Historically, cereal crops have beenimportant in providing food and nesting habitat but lesscommonly used now. Will search hedges and headlandsfor seeds, buds and fungal material. Field headlands,rough grass banks and hedgerows act as refuge andreservoir during winter.
Associated crops: Cereals, legumes, long grass, especially traditional hayfields.
Recorded in: 693 10 km squares in Britain
Population: Declining
( 6 - 8Body weight (g)
Foragingbehaviour:
Active both day and night, especially in summer withcrepuscular peaks of activity. Mean home ranges approx.350 - 400 sq m. Acute hearing and good sight. Will takecereal grains directly from cereal head.
( 24
Outline diet: In the field, takes buds in spring and grain from cerealheads in summer. Urban populations will take seeds, fruit,leaves (monocot and dicot), insects, fungi, moss and rootmaterial. In captivity will eat hard and soft bodied insectsas well as fruits, seeds and berries.
Status: Locally common
%)
Density: Dramatically variable. Populations can peak at over 200per ha. one year followed by several years of apparentdecline. Between July and October, mean densities perha of 0.05 in barley, 0.4 in wheat, 2.5-5.0 in roughmeadow and 20 in reedbeds.
)
England:
1,415,000
Scotland:
0
Wales:
10,000
Trend:
26
Rodentia Muridae
Harvest mouse Micromys minutus
Further notes on use made of arable land:
The harvest mouse was the rodent species least often encountered in theBoxworth study (Johnson et al, 1992). Because of its climbing habit in thesummer, traps laid on the ground may underestimate true numbers. Signs ofharvest mouse activity were recorded in all three study fields.
Modern agricultural practice appears to have led to a decline in numbers.Combine harvesting may kill mice nesting in cereal fields and unthreshed cornricks were historically an important winter habitat which exixt no longer (Tew,1994).
27
Insectivora Erinaceidae
Hedgehog Erinacus europaeus
1100
Young born from: May to: Sep
Geography: Throughout mainland Britain but scarce in wet habitats,large arable fields and conifer plantations. Less abundantin the south.
Feeding habitat: Feed almost wholly on invertebrates found on ground.Prey is most abundant where there is close proximity ofgrassland to woodland. Also common in suburban areaswhere suitable cover exists.
Associated crops: Pasture, grassland.
Recorded in: 1993 10 km squares in Britain
Population: Slight decline
( - 1600Body weight (g)
Foragingbehaviour:
Solitary, nocturnal. Food caught and killed entirely bymouth. Poor eyesight but good hearing and smell. Snuffleaudibly when searching for food on surface. Do notnormally dig. Will range 0.5 - 1.5 km in a night covering 2-5 ha. Adult males may travel up to 3 km over 12 ha. Nocaching of food for winter.
( 70
Outline diet: Ground-living invertebrates such as earthworms, slugs,beetles, caterpillars, fly larvae, centipedes and spiders.Small numbers of birds' eggs, chicks and carrion alsotaken.
Status: Common
%)
Density: Densities probably lower in rural than suburban areaswhere they may reach 1 per ha. Optimal rural habitat 1per 2.5 ha and 1 per 20 ha for forestry plantations,improved grassland and arable areas
)
England:
1,100,000
Scotland:
310,000
Wales:
145,000
Trend:
28
Insectivora Erinaceidae
Hedgehog Erinacus europaeus
Further notes on use made of arable land:
There is little information on hedghogs' use of arable land but a radio-trackingstudy at 3 permanent pasture sites in Oxfordshire indicated an average homerange of 2324 sq m. The overlap between neighbouring home ranges wassmall -- 12.8%.
29
Perissodactyla Equidae
Horse Equus caballus
Young born from: Apr to: Sep
Geography: Free ranging populations found in New Forest, Dartmoor,Exmoor, the Gower peninsula, Lake district,Northumberland, Shetland, certain Hebridean islands andin W Ireland.
Feeding habitat: Marginal habitats such as open moorland, roughgrassland with access to woodland cover. Strongpreference for grassland during summer and woodlandduring winter.
Associated crops: Rough grazing and marginal areas.
Population: Stable
( 200000 - 250000Body weight (g)
Foragingbehaviour:
Preferential grazers during summer, being relativelyunselective bulk feeders. As grass productivity decreasesthrough summer, increased browsing occurs.
Outline diet: Unselective grazer. During summer, diet mostly consistsof grass. As grass becomes scarcer in winter,consumption of gorse, tree leaves, moss and heatherincreases.
Density: Controlled by man.
)
England: Scotland: Wales: Trend:
30
Carnivora Mustelidae
Mink Mustela vison
Young born from: May to: May
Geography: Widespread in mainland Britain and Ireland, althoughsome areas remain incompletely colonised;- N and WIreland, NW tip of Wales, N of Scotland and either side ofthe Wash, and E Yorkshire.
Feeding habitat: Wide range of aquatic habitats, preferring eutrophicstreams and rivers which have abundant bank side cover.Will spend time away from water habitat hunting for prey,for example where rabbits plentiful.
Recorded in: 1051 10 km squares in Britain
Population: Increasing
Male: 1150
Female: 650 ( 437 -
( 840 - 1805 )Body weight (g)
Foragingbehaviour:
Predominately nocturnal but can be active at any time.Regular patrolling patterns observed. Activity ofteninfluenced by that of prey. Home ranges linear and 1 - 6km in length. Swim well, chasing and catching fishunderwater. Prey killed with bite to back of neck. Will takebirds, small mammals, fish, carrion and very commonlyrabbits.
( 37
Outline diet: Opportunist predators. Fish comprise single mostimportant food source but mammals (especially rabbitsand water voles) and birds (especially ducks, moorhensand coots) are also important.
Status: Increasingly widespread
%)
Density: In Britain, highest recorded density is one territory per 0.5km of rocky shore. In a Devon river environment, meandensity was 0.88 per km river.
810 )
England:
46,750
Scotland:
52,250
Wales:
9,750
Trend:
31
Insectivora Talpidae
Mole Talpa europea
Young born from: Apr to: Jul
Geography: Throughout mainland Britain, Anglesey, Isle of Wight anda few Inner Hebrides. Absent from Ireland.
Feeding habitat: Wherever the soil is deep enough to tunnel. Origins are indeciduous woodland but abundant in pasture and arableland. Uncommon in coniferous woods, moorland or sand-dunes. Move into arable l0and from field boundaries afterploughing.
Associated crops: Pasture, arable land.
Recorded in: 2225 10 km squares in Britain
Population:
Male: 110
Female: 85 ( 72 -
( 87 - 128 )Body weight (g)
Foragingbehaviour:
Food items obtained almost exclusively in tunnel whichacts as pitfall for soil invertebrates. Incidental food takenwhilst burrowing. Rarely seen on surface. Solitary for mostof year; female home range 1000-2000 sq m. Male rangecan be 8000 sq m during mating period. Both sexes have3 periods of activity per day each of 3-4 hours durationfollowed by similar rest periods. During autumn only 2periods of activity for males. Moles will often clean wormsby squeezing them through their fingers as they eat,forcing most of the soil from the gut. This may reduceintake of soil contaminants such as pesticide residues.
( 78
Outline diet: Earthworms, especially Lumbricus terrestris are singlemost important prey. Insect larvae taken in largenumbers, myriapods and molluscs less so. Diet reflectsprey abundance; worms more important in winter; insectspredominate in summer.
Status: Common
%)
Density: Poor habitat 1.3 per ha. Good habitat 4 per ha. Densitiesof 8 per ha. in winter and 16 per ha. in summer recordedin English pastures. In Scotland, densities in woodlandand pasture remain similar throughout year at 4-5 per ha.
106 )
England:
19,750,000
Scotland:
8,000,000
Wales:
3,250,000
32
Artiodactyla Cervidae
Muntjac Muntiacus reevesi
Young born from: Jan to: Dec
Geography: Well established in S England from E Anglia up toMidlands and west to Wales and Devon.
Feeding habitat: Previously thought to prefer dense habitat with diversity ofvegetation. Now thought to be more cosmopolitan(Chapman et al, 1994). Frequently present in commercialconiferous plantations where deciduous trees are presentand a variety of shrubs and ground cover can be found.Also found in suburban habitats where over-growngardens are present.
Associated crops: Mixed forestry plantations
Recorded in: 417 10 km squares in Britain
Population: Increasing
Male: 14800
Female: 12200 ( 9000 -
( 10500 - 18300 )Body weight (g)
Foragingbehaviour:
Main activity and feeding periods are crepuscular and inthe middle of the day. Basically solitary with home rangesc. 14 ha. in varied deciduous woodland. Prefers to browse.
( 15
Outline diet: A browser rather than a grazer. Brambles important allyear round. Other food items change seasonally includingivy, ferns, fungi, broad leaves, shrubs, nuts and otherfruit. Grasses are significant in the spring. Rose,dogwood, blackthorn and hawthorn may be important inthe summer.
Status: Locally common
%)
Density: Variable with habitat; mixed woodland can support oneMuntjac per 6.8 ha although can be higher in more variedthicket woodland.
15800 )
England:
40,000
Scotland:
50
Wales:
250
Trend:
33
Artiodactyla Cervidae
Muntjac Muntiacus reevesi
Further notes on use made of arable land:
Although largely preferring to forage in woodland, Muntjac at Monks WoodNature Reserve were increasingly observed in surrounding arable land duringFebruary and March, when woodland food resources were scarce (Cooke, 1996).
34
Carnivora Mustelidae
Otter Lutra lutra
Young born from: Jan to: Dec
Geography: Throughout Ireland and much of Scotland althoughnumbers much reduced in areas of intense agriculture.Present in Wales (except South) and in SW England andfar East, but absent from central England
Feeding habitat: Lakes, rivers, streams and marshes. Capable of travellingoverland between watersheds and of living in marinehabitats as well as in freshwater.
Recorded in: 1308 10 km squares in Britain
Population: Slight increase
Male: 10100
Female: 7000
Body weight (g)
Foragingbehaviour:
On rivers, predominately nocturnal and crepuscular.Female otters have recorded ranges of 16 - 22 km, malesgreater. In clear, well lit water, otters hunt by sight; snoutvibrissae used to detect fish movements when visibility ispoor.
( 46
Outline diet: Mainly fish but opportunistic and will take waterfowl, frogsand crayfish. Rarely take mammals but diet may includerabbits and water voles if common. Aquatic insects areoccasionally taken.
Status: Endangered
%)
Density: Difficult to assess. In Perthshire, 4 adults occupied 57.4sq km, while in E. Anglia 3 re-introduced adults occupied74.7 sq km.
England:
350
Scotland:
6,600
Wales:
400
Trend:
35
Chiroptera Vespertilionidae
Pipistrelle Pipistrellus pipistrellus
Young born from: Jun to: Jul
Geography: Widespread through most of Britain and Ireland. Notresident in Shetland
Feeding habitat: Found in a wide variety of habitats including farmland,open woodland, suburban gardens, lakes and rivers.Avoid very open country. Will often roost in buildings,rarely in caves.
Recorded in: 1441 10 km squares in Britain
Population: Declining
Male:
Female: ( 5.15 -
( 4.2 - 5.7 )Body weight (g)
Foragingbehaviour:
Patrol in regular beats where insect densities aregreatest -- along linear vegetative features, around treesand low over water. Feeding occurs mostly around duskand dawn in response to insect density. Feeding ratesmay reach 20 captures a minute.
( 51
Outline diet: Unselective feeders taking flying insects in proportion totheir availability. Bulk of diet consists of midges, gnatsand mosquitoes.
Status: Common and widespread
%)
Density: Minimum of 0.05 breeding females per ha. Population in1987 estimated as having declined to 38% of thatrecorded in 1978.
7.05 )
England:
1,250,000
Scotland:
550,000
Wales:
200,000
Trend:
36
Carnivora Mustelidae
Polecat Mustela putorius
Young born from: May to: Jul
Geography: Currently restricted to Wales and adjacent Englishcounties only. Formerly widespread in British Isles butnever present in Ireland.
Feeding habitat: Woodland, farmland, riverbanks and around farmbuildings where there exists cover and prey (rats andmice). Also attracted to rubbish tips.
Recorded in: 235 10 km squares in Britain
Population: Slight increase
Male: 1111
Female: 689 ( 530 -
( 800 - 1710 )Body weight (g)
Foragingbehaviour:
Mainly nocturnal and crepuscular in summer and diurnal inwinter. Typically covers 3 - 4 km per night. Very keensense of smell and hearing. Prey killed by bite to back ofneck. Some food taken as carrion, also birds eggs taken.Heads of toads not eaten because of poison glands; frogsovary masses also rejected. Will cache food if abundant.
( 8
Outline diet: Principally rabbits but will also take small rodents, birdsand amphibians. No evidence in the UK of fish or plantmaterial in the diet
Status: Locally common
%)
Density: Exact figures not known, but assumed quite low. Swissdata suggest densities of 0.1 -- 1 per sq km.
915 )
England:
2,500
Scotland:
0
Wales:
12,500
Trend:
37
Insectivora Soricidae
Pygmy shrew Sorex minutus
4
Young born from: Apr to: Oct
Geography: Throughout Britain, including the larger islands andthroughout Ireland (unlike common shrew). Widespread inat all altitudes
Feeding habitat: Preference for grassland but like the common shrew, theyare widespread wherever there is ground cover. Lessnumerous than common shrew in lowland habitataccounting for 12% of captures on woodland and 10-18%on grassland.
Associated crops: Rough grazing, permanent pasture, set-aside, agriculturalheadlands.
Recorded in: 958 10 km squares in Britain
Population:
( 2.4 - 6.1Body weight (g)
Foragingbehaviour:
Unlike the common shrew, bulk of diet is found on thesurface (60%) rather than underground (10%). Ratherweak burrowers though they will make use of naturalcrevices and other animal burrows. More active duringdaytime than common shrew but equally active at night.Main peaks of activity between 0800-1000 and 2100-2300hours.
( 34
Outline diet: Spiders, harvestmen, beetles and woodlice are mostcommon prey items in England.. Opportunist predators.Prey taken in proportion to their availability, typicallybetween 2-6mm in length. Do not take earthworms butwill occasionally eat small slugs, snails and plant material.Reject millipedes.
Status: Common and widespread
%)
Density: Except for moorland, densities much less than commonshrew. Mean of 6, maximum of 12 per hectare ongrassland.
)
England:
4,800,000
Scotland:
2,300,000
Wales:
1,500,000
38
Insectivora Soricidae
Pygmy shrew Sorex minutus
Further notes on use made of arable land:
Tew et al (1994a,b), found that both common and pygmy shrews were caughtentirely in the hedgerows during the winter months, but as crop cover increasedover the summer they were occasionally caught in the cereal field itself, thoughrarely more than 20m from a hedgerow.
39
Lagomorpha Leporidae
Rabbit Oryctolagus cuniculus
1500
Young born from: Feb to: Sep
Geography: Widespread in Britain and Ireland up to tree line; on mostsmall islands except Isles of Scilly. Most abundant in Sand E England.
Feeding habitat: Areas of short grass; naturally occurring, dry heaths orclosely grazed agricultural pastures with secure refugenearby. Never abundant in large coniferous plantations.
Associated crops: Pasture, newly sown rye grass and cereals, root crops,horticulture, new forestry plantations.
Recorded in: 2267 10 km squares in Britain
Population: Increasing
( 1200 - 2000Body weight (g)
Foragingbehaviour:
Activity mainly nocturnal and crepuscular. Home rangesare small, 0.3 - 3 ha. Territories occupied as a social group.
( 80
Outline diet: A wide range of herbs and grasses, preferring succulentleaves and shoots and selecting more nutritious specieseg Festuca grasses. Bark of trees eaten especially duringsnow cover.
Status: Abundant and widespread
%)
Density: Seasonal; 1 - 15 per ha. in winter; 1 - 40 during summerpeaks. Populations highest on sandy soils. JNCCestimates are: 5 per ha for optimal habitat, 2.5 per ha forlowland grassland and arable land.
)
England:
24,500,000
Scotland:
9,500,000
Wales:
3,500,000
Trend:
40
Lagomorpha Leporidae
Rabbit Oryctolagus cuniculus
Further notes on use made of arable land:
Rabbits were observed over 2 winters foraging on and around a winter barleyfield. Intensity of feeding declined exponentially with distance from the field edgeand cover. Only 5% of rabbit sightings were at distances greater then 40m fromthe field edge. Percentage of time spent feeding was greatest between 5 and30m metres into the crop. Individuals spent more time in vigilance behaviour atthe extreme edge and towards the centre of the field. Although some rabbitswere active on the field throughout the night, on average only a third of therabbits using the field were present at any one time. The probability of feeding inthe field was related to the location of an individual's daytime refuge. If theanimal had to travel more than 100m further to feed at the study field comparedwith an alternative site, then it would only rarely appear at the study field(Cowan, 1989).
41
Rodentia Sciuridae
Red squirrel Sciurus vulgaris
300
Young born from: Feb to: Aug
Geography: 10 isolated populations within Britain; including Brownseaisland, Snowdonia, Lake district, Epping forest and Isle ofWight. Majority of Ireland and Scotland except West coastand very North, respectively.
Feeding habitat: In Britain populations most stable in large tracts ofconiferous woodland (>100 ha.) over 25 years of age.Scots pine, Norway spruce and Siberian pine forestpreferred here. Also found in smaller woods where Greysquirrel absent.
Associated crops: Coniferous woodland.
Recorded in: 386 10 km squares in Britain
Population: Declining
( 220 - 440Body weight (g)
Foragingbehaviour:
Diurnal. Home range highly variable around 7.4 ha.; largerin deciduous than coniferous forests. Solitary butcommunal nesting can occur in winter. Very arboreal(unlike grey squirrels) spending 70% of their time off theground. Active all year so will cache food in latesummer/autumn. Very good sense of smell and hearingand has wide-angled vision for foraging.
( 14
Outline diet: Mainly tree seeds, hazelnuts, beech mast, acorns, conifercones and fungal fruiting bodies, but will also take buds,shoots and flowers. Bark eaten at any time of year. Othergreen material, invertebrates and lichen may also beeaten. Less able, than grew squirrel to exploit acorncrops.
Status: Localised
%)
Density: Long term average densities of 0.5 - 1.5 per ha. Numberslowest in spring, highest in autumn.
)
England:
30,000
Scotland:
121,000
Wales:
10,000
Trend:
42
Artiodactyla Cervidae
Roe deer Capreolus capreolus
Young born from: May to: Jul
Geography: Widespread throughout Scotland and N England. Presentthroughout most of S and England. Absent from Ireland.
Feeding habitat: Open mixed, coniferous or deciduous woodland. Also willoccupy agricultural land if small woods are available forcover.
Associated crops: Forestry, oilseed rape, wheat, barley, maize, root crops.
Recorded in: 1237 10 km squares in Britain
Population: Increasing
Male: 23900
Female: 22300 ( 18000 -
( 18000 - 28500 )Body weight (g)
Foragingbehaviour:
Generally 9 periods of activity throughout 24 hours withlargest feeding bouts at dusk and dawn. Generally ventureonto agricultural land during night. Territorial during springand summer but will form groups of 8-12 to forage duringwinter. Well developed senses of smell and hearing, butsight more suited to detecting movement.
( 43
Outline diet: Opportunists. Diet varies according to season andavailability. Deciduous buds and shoots, non grass herbsand shrubs (especially brambles) important throughoutthe year. Ivy, conifers and dwarf shrubs taken in winter.Small amounts of grasses eaten throughout the year.Cereal crops eaten in early stages of rapid growth andagain as they ripen.
Status: Common
%)
Density: Plantation age and structure influences local density,highest (>25 per sq km) in young plantations 5-15 yearsold, decreasing to c.8 deer per sq km in 30-45 year oldstands.
28000 )
England:
150,000
Scotland:
350,000
Wales:
50
Trend:
43
Artiodactyla Cervidae
Roe deer Capreolus capreolus
Further notes on use made of arable land:
A study of nocturnal feeding habits in Scotland in January to March showed thatgrass, stubble and ploughed fields were all strongly avoided, while winter barleywas preferred in all 3 months. Oilseed rape was preferred in January andsubsequently avoided. The majority of deer were seen within 75m of woodlandedge (Boag et al, 1990).
Deer were observed along standard transects on mixed arable farmland andwoodland in Hampshire. Deer were seen feeding on kale and fodder rootsduring autumn and on cereals between March and early June. Pastures wereused extensively in spring and early summer. Woodland was an importanthabitat all the year round (Putman, 1986).
A Polish study found that Roe deer density was at it's highest in small woodssurrounded by agricultural land. An analysis of density of faecal pellets,supplemented by visual sightings showed that two thirds of feeding timeoccurred within 200m of the forest edge, with individuals only rarely penetratingmore than 500m into surrounding arable landscape (Aulak & Babinska-Werka,1990). A radio-tracking study in northern France (Cibien, 1995) also indicatedthat the majority of locations were in non-cultivated areas but that analysis ofrumen contents revealed a high dependence on cultivated plants, especially inwinter.
44
Carnivora Mustelidae
Stoat Mustela erminea
Young born from: Apr to: May
Geography: Throughout mainland Britain and Ireland at all altitudes.
Feeding habitat: Can be found in wide range of habitats that offer foodsource and cover, farmland, woodland, moors andmarshes. Tend to prefer early successional vegetationwhich is more favoured by prey.
Recorded in: 1519 10 km squares in Britain
Population: Slight decline
Male:
Female: ( 190 -
( 260 - 320 )Body weight (g)
Foragingbehaviour:
Nocturnal in winter, diurnal in summer. Spells of activity10 - 45 minutes in length, interrupted by periods of rest.Always active when foraging, searching along hedges andwalls concentrating on most likely prey hideouts. Preylocated by movement and sound. Kill made by bite to backof neck. Will enter underground tunnels in search of watervoles and rabbits. Will avoid eating fur, which is used toline den.
( 53
Outline diet: Mainly small mammals (rabbits, water voles, rats), birdsand eggs, depending on what is available.
Status: Common
%)
Density: Density very much dependant upon prey (rabbit). InSweden, <1 - 2.2 stoats per 10 sq km, depending onhabitat. Autumn densities from Europe and Canadaaverage 3- 10 Stoats per sq km.
220 )
England:
245,000
Scotland:
180,000
Wales:
37,000
Trend:
45
Insectivora Soricidae
Water shrew Neomys fodiens
Young born from: May to: Sep
Geography: Throughout mainland Britain but rather patchy in Scotlandand its islands. Absent from Ireland.
Feeding habitat: River banks, preferably where water is fast flowing andunpolluted. Also occur by ponds and drainage ditchesand, in NW Scotland, on rocky beaches. Will usehedgerows on arable land as corridors between morefavoured habitat.
Associated crops: Watercress beds, riparian margins.
Recorded in: 654 10 km squares in Britain
Population: Slight decline
( 12 - 18Body weight (g)
Foragingbehaviour:
Forage in riparian habitat and underwater. Active day andnight with peak activity prior to dawn. Daily movementapprox. 10 - 60 m, rarely 150 - 200 m. Nomadic existence,moving home ranges after a few months. Very goodswimmers, ability to dive well. Sensitive, mobile vibrissaeon snout may assist in prey detection. Well developedhearing and smell. Saliva is venomous. Aquatic preybrought ashore to be eaten. Food caches sometimesmade.
( 23
Outline diet: Will forage underwater and on land for a wide variety ofinvertebrates, small fish and amphibians. Dominantfreshwater prey in all months include freshwatercrustaceans, (Gammarus and Asellus) and cased Caddisfly larvae. Other major prey items include terrestrialbeetles, spiders, centipedes, molluscs and earthworms.
Status: Locally common
%)
Density: Much less prolific than Common Shrew. Estimate of 3.2per ha. in English Water Cress beds but probably under-estimate.
)
England:
1,200,000
Scotland:
400,000
Wales:
300,000
Trend:
46
Rodentia Muridae
Water vole Arvicola terrestris
Young born from: Apr to: Sep
Geography: Throughout England, Wales and south Scotland, generallyconfined to low ground near water. Very local populationsin NW and N Scotland. Absent from Ireland.
Feeding habitat: Densely vegetated banks of ditches, rivers and streams,where the current is slow and water present year round.Less frequently found on ponds. Can live away from water.
Recorded in: 1060 10 km squares in Britain
Population: Declining
Male: 311
Female: 272 ( 225 -
( 246 - 386 )Body weight (g)
Foragingbehaviour:
Active every 2 - 4 hours more throughout day than night.In winter, activity outside burrow is rare. Females haveexclusive territories, males staying with females for entireseason. Voles swim frequently, diving well. Visual cuesare particularly important, water voles showing an ability toremember changes to foraging routes.
( 37
Outline diet: Largely vegetarian taking reed grasses (phragmites) andsedges, sometimes rushes and dicotyledons such asnettles.
Status: Common but declining
%)
Density: Numbers per km of riverbank vary from 17 in South Westwater region to 43 in Anglian and Southern regions.Densities will generally be lower and will vary with bankcondition.
310 )
England:
752,000
Scotland:
376,000
Wales:
41,000
Trend:
47
Carnivora Mustelidae
Weasel Mustela nivalis
Young born from: Apr to: Sep
Geography: Throughout British mainland and most islands except Isleof Man and most Hebrides. Absent from Ireland
Feeding habitat: Very adaptable, yet restricted mainly to hedgerows, dykesand other forms of cover where rodent food items aremost available. Very successful in searching for food insnow in winter.
Recorded in: 1490 10 km squares in Britain
Population: Slight decline
Male: 120
Female: 63 ( 55 -
( 106 - 131 )Body weight (g)
Foragingbehaviour:
Active both day and night for bouts of 10 - 45 minutesinterrupted by periods of rest. Weasels will enter tunnels ofvoles and mice but not those of rabbits. Prey located bysight, smell or sound. Lethal bite to back of neck. Willavoid eating fur, which they use to line den. Will cacheprey when abundant.
( 52
Outline diet: Specialise on small rodents (especially voles) but will alsotake birds and eggs when available. Wood mice takenmost often after harvest and young rabbits may be takenin the spring. Males tend to take larger prey (rats,lagomorphs) than females. Both sexes will raid birdnestboxes.
Status: Common
%)
Density: Very variable and closely related to fluctuations in rodentpopulations. In English woodland densities can vary from1 - 4.5 weasels per ha.
69 )
England:
308,000
Scotland:
106,000
Wales:
36,000
Trend:
48
Rodentia Muridae
Wood mouse Apodemus sylvaticus
18
Young born from: Apr to: Oct
Geography: Ubiquitous in Britain and Ireland except on most openmountainsides
Feeding habitat: Ground dwelling amongst thick grass, woodland, arableland, heather and sand dunes. Like cover offered by drystone walls, hedgerows and gardens and even buildings ifunoccupied.
Associated crops: Set-aside, arable crops, especially cereals, sugar beet andoilseed rape.
Recorded in: 1379 10 km squares in Britain
Population: Stable
( 13 - 27Body weight (g)
Foragingbehaviour:
Nocturnal behaviour. Newly planted sugar beet seedsdetected by olfaction, dug up, coating removed, pericarpcracked and embryo eaten. Will climb hedges in autumnand winter for fruits, esp rose hips. Summer home rangesare about 2000 sq m in deciduous woodland, with highervalues in less productive habitats. Communal nestingobserved in winter.
( 48
Outline diet: Opportunists, taking mainly seeds and invertebrates. Incereal fields will feed on grain shed after harvest,arthropods in spring and, weed and grass seed insummer. Sown seed is also taken. In sugar-beet fields,weed seeds taken before autumn harvest, with insectlarvae, worms and beet remains in winter.
Status: Common
%)
Density: Variable. Densities of 1 - 40 per ha. in mixed deciduouswoodland in summer can increase to 130 - 200 per ha. inwinter if there was a good autumn seed crop. On arableland densities range from 0.5 per ha in summer to 17.5per ha in winter.
)
England:
19,500,000
Scotland:
15,000,000
Wales:
3,500,000
Trend:
49
Rodentia Muridae
Wood mouse Apodemus sylvaticus
Further notes on use made of arable land:
The typical use of arable land by the wood mouse is described by Tew (1994) as follows: " thewood mouse is found on open arable land throughout the year. However, the suitability of arablefields for small mammals is seasonally variable and hedgerows are valuable resources for woodmice through the winter, providing both food and cover, although the open field is still exploited bysome sections of the population. As the cover and food abundance in the fields increases duringspring and summer, mice move out into the fields, nesting, mating and foraging entirely within thecrop."
McDonald et al (unpublished data) studied wood mice in and around Oxfordshire wheat, barleyand oilseed rape fields. Fewer mice were trapped in rape fields than in barley or wheat fields.Radio-collared mice were also found to move more quickly though rape than through the othercrops. The authors speculate that rape may have fewer weed patches known to be attractive towood mice. An earlier study by Green (1979) on mice inhabiting open arable land with fewhedges, found little marked preference between crops in winter but a marked preference forwinter wheat fields in spring and early simmer.
Reduced application of herbicides in small experimental conservation plots along the edge ofcereal fields led to increased weeds and invertebrates. Radio-collared wood mice sought out suchareas for feeding (Tew et al, 1992). On a larger scale the Boxworth project failed to show anyconsistent effect of pesticide treatment on trapping success (Johnson et al, 1992). On a Scottishfarm consisting of a mosaic of crops, permanent set-aside and semi-natural vegetation, radio-collared wood mice showed no clear habitat preferences, nesting and foraging in all habitat types(Rogers, 1993). Season and agricultural practice had no effect on preference, although woodmice in Scottish set-aside had smaller home ranges than those in English arable land. Theauthors speculate that food was more abundant at the Scottish sites. (Rogers & Gorman, 1995).Tattershall et al (1997) found summer trapping success to be significantly lower in set-aside(0.6%) than in cereal crops (13%). There was no evidence that wood mice preferred strips toblocks of 1-yr set-aside or that the animals distinguished between sown and naturally regeneratedset-aside, although the authors note that cover was thin in all cases.
Radio-collared wood mice dosed with dimethoate and released into cereal fields were less activefor the first 6 hours but thereafter appeared to recover, and medium term survival was notaffected (Dell'Omo & Shore, 1996). Methiocarb slug pellets broadcast in arable fields in autumnand spring led to decreases in the wood mouse population of 78% and 33% respectively (Shoreet al, 1997). Wood mice appear to be particularly vulnerable to seed treatments. Sowing withseeds treated with bendiocarb resulted in a population reduction and 46% of the individualscaught had residues detectable in the gut (Tarrant et al, 1990).
Wood mice living in cereal fields were not directly effected by harvesting operations but thesubsequent loss of cover had important consequences. More than half of the radio-collaredindividuals were taken by predators in the week following harvest. Other individuals eitheremigrated from the fields or reduced their activity (Tew et al, 1993). In southern Sweden, Loman(1991) notes that wood mice burrows are not destroyed by ploughing and wood mice are found incropped fields throughout winter.
Wood mouse populations in woodland are strongly influenced by the neighbouring croppingpattern. In spring, mice were less abundant in those woods with more wheat and rape fieldsaround them (Fitzgibbon, 1997).
Wood mice in autumn in hedgerows show a strong association with berry abundance (Poulton,1994). Woods et al., (1996) found that wood mice were more likely to be trapped in newly plantedhedgerows than in older, established ones.
An analysis of barn owl pellets at 15 sites in Haute Savoie, France, suggested a strong positiverelationship between the percentage occurrence of wood mice and the total area under cerealcultivation (Taberlet, 1986).
50
51
Detailed Dietary Information ( 30 species in alphabetical order)
from till food type g/day kJ/day % of diet referencecomment
Badger Meles meles
Aug Aug Earthworms Barker, 1969aUnpublished data from a wet summer. Wiytham wood,Oxford. Quoted in Neal, 1986. (Method of analysis notspecified.)
40
Wheat 30
Aug Aug Wheat Barker, 1969bData from a dry summer75
Insects 15
Fruits Biancardi et al., 1993Faecal analysis. Italy. Fruits single most important foodtype, especially chestnuts. Cereals, grass and roots,birds, mammals and gastropods represent 10% of diet.
No seasonal data
Earthworms 642 Neal, 1986642g of earthworms found in the stomach of a female inApril in Somerset. Several other stomachs containedmore than 200 earthworms, a volume of more than1000ml.
No seasonal data
Jan May Insects Pigozzi, 1988% VOLUME in faeces. Combined volume of fruit andinsects in diet was greater than 89% of total food eatenover 2 years in Italy where earthworms are notcommon. Insects eaten between late winter and earlysummer.
45
Aug Dec Fruit Fruits mainly eaten late summer to midwinter.44
52
from till food type g/day kJ/day % of diet referencecomment
Badger Meles meles
Animal Roper & Lups, 1995% VOLUME in 217 stomach contents. SwitzerlandOverall, faunal material consisted 55% diet,Earthworms most common but not eaten in largevolumes. Voles and insects eaten most of year butnever in large quantities, whilst wasps were eatenseasonally and in large volumes. Total food intake isgreatest in autumn, while diversity is greatest insummer.
No seasonal data 55
Vegetal Maize eaten most of the year but not in large quantities.Cherries, plums and oats eaten seasonally in largequantities.
45
Animal Roper & Mickevicius,1995
% VOLUME in stomach or faeces. Review of 69 studiesfrom the Soviet Union.The most important animal foods were insects (30%)and small mammals (20%). Earthworms neverexceeded 5%. Other animal foods included birds,reptiles, amphibians, fish, molluscs and carrion.
No seasonal data 62
Vegetal Fruits seasonally important; roots, tubers, bulbs, cerealsand nuts also eaten.
38
Insects 30
Small mammals 20
Sep Nov Grapes Roper et al, 1990% VOLUME in faeces from territories known to overlapvineyards in E. Sussex.Three radio-collared individuals were observed to spend30min to 2h in the vineyard each night.
64
53
from till food type g/day kJ/day % of diet referencecomment
Badger Meles meles
Earthworms Roper, 1994In a review of 11 quantitative studies, no study showedearthworms to comprise more than 50% of seasonallyadjusted diet. In 2 studies fruit and insects accountedfor 95 - 97% of total consumption.
No seasonal data 50
Wheat & Barley Roper, et al., 1995Observational study on South Downs, UK.Wheat preferred to Barley. Wheat taken from early May(milk stage) to November. Radio-collared individualsspent more than 30% of foraging time in wheat fields.Barley taken between July and September, but only bysome individuals.Some individuals may consume as much as 1100mlgrain in a night although 50-250ml is more common.
No seasonal data
Jan Feb Earthworms Sheperdson et al.,1990
%VOLUME in faeces. Mixed farming area in Sussex.Diet dominated by earthworms in winter and spring andby cereals and fruit in summer and autumn.Radiotracking of 25 individuals revealed a preferencefor pasture in the spring which declined through the yearin favour of wheat and other arable land.
60
Grass 25
Mar Apr Earthworms 50
Grass 20
Insects 5
Wheat 5
May Jun Wheat 45
Insects 15
Earthworms 10
54
from till food type g/day kJ/day % of diet referencecomment
Badger Meles meles
May Jun Grass Sheperdson et al.,1990
10
Jul Aug Wheat 70
Insects 10
Earthworms 5
Grass 5
Sep Oct Wheat 50
Fruit 25
Earthworms 10
Nov Dec Earthworms 60
Fruit 15
Wheat 10
Mar May Earthworms Skinner & Skinner,1988
% VOLUME in 315 Faecal samples. Population of 15adults in 41 sq km arable land in Essex, UK. Suboptimalhabitat for badgers
33
Beetles 20
Noctuid larvae 13
Mammals 11
Birds 10
Wheat seed 3.6
Jun Aug Wheat seed Wheat ears often taken green.62
Beetles 14
Earthworms 8
55
from till food type g/day kJ/day % of diet referencecomment
Badger Meles meles
Jun Aug Bumble bees Skinner & Skinner,1988
7
Sep Nov Wheat seed Will glean wheatfields after stubble has been burnt.28
Fruit 13
Beetles 12
Earthworms 10
Other crops Mostly maize left for pheasant cover10
Tree seeds Predominantly acorns10
Noctuid larvae 8
Dec Feb Wheat seed Possibly rooted from ground, scavenged from grainstores or from pheasant feeders
35
Tree seeds 29
Earthworms 14
Beetles 8
56
from till food type g/day kJ/day % of diet referencecomment
Bank vole Clethrionomys glareolus
Plants & fungi Castien & Gosalbez,1996
Analysis of stomach contents in the Western Pyrenees.Food intake mainly leaves, although fungi becomeimportant in summer and autumn. Seeds unimportant.Carnivory low.
No seasonal data
Jan Ash seeds Flowerdew &Gardner, 1978
% VOLUME in stomach contents. Derbyshire woodland.UK.Ash seeds eaten throughout the year but highlyseasonal, reaching a peak (84%) in January and atrough (3%) in July.
84
Galaeobdolonstems
12
Fungus on wood 3
Apr Galaeobdolonleaves
36
Ash seeds 36
Galaeobdolonstem
12
Bark 10
Jul Ash leaves (live) 52
Ash stems 21
Ash leaves (dead) 4
Ash seeds 3
Sep Fruits Mainly hawthorn berries32
Barley testa &endosperm
27
57
from till food type g/day kJ/day % of diet referencecomment
Bank vole Clethrionomys glareolus
Sep Brachipodiumtesta &endosperm
Flowerdew &Gardner, 1978
19
Ash seeds 16
Oct Barley testa &endosperm
32
Ash leaves (dead) 16
Ash leaves (live) 13
Ash seed 11
Mar Mar Seeds Hansson, 1971% NUMBER of items found in coecae of bank volescaught in Southern Swedish forest.N animals = 30, % OCCURRENCE = 57%
30
Insects, adults N= 30, % OCCURRENCE = 40%21
Fungi N= 30,. % OCCURRENCE = 20%11
Moss N= 30, % OCCURRENCE = 2011
Insects, larvae N= 30,. % OCCURRENCE = 17%9
Herbs N= 30, % OCCURRENCE = 13%7
Grass N= 30, % OCCURRENCE = 10%5
Lignified stems N= 30, % OCCURRENCE = 3%2
Apr May Insects, adults N= 66, % OCCURRENCE = 44%19
Herbs N= 66, % OCCURRENCE = 39%17
Seeds N= 66, % OCCURRENCE = 29%13
Moss N= 66, % OCCURRENCE = 24%11
Fungi N= 66, % OCCURRENCE = 23%10
58
from till food type g/day kJ/day % of diet referencecomment
Bank vole Clethrionomys glareolus
Apr May Insects, larvae Hansson, 1971N= 66, % OCCURRENCE = 20%9
Lignified stems N= 66, % OCCURRENCE = 14%6
Lumbricids N= 66, % OCCURRENCE = 11%5
Grass N= 66, % OCCURRENCE = 9%4
Molluscs N= 66, % OCCURRENCE = 2%1
Graminoids N= 66, % OCCURRENCE = 2%1
Ferns N= 66, % OCCURRENCE = 2%1
Roots N= 66, % OCCURRENCE = 3%1
Jun Aug Insects, adults N= 87, % OCCURRENCE = 67%26
Herbs N= 87, % OCCURRENCE = 59%23
Seeds N= 87 % OCCURRENCE = 32%12
Lumbricids N= 87. % OCCURRENCE = 23%9
Moss N= 87, % OCCURRENCE = 20%8
Insects, larvae N= 87, % OCCURRENCE = 17%7
Grass N= 87, % OCCURRENCE = 13%5
Fungi N= 87, % OCCURRENCE = 13%5
Lignified stems N= 87, % OCCURRENCE = 12%4
Berries N= 87. % OCCURRENCE = 2%1
Sep Nov Seeds N= 108, % OCCURRENCE = 59%22
Insects, adults N= 108, % OCCURRENCE = 41%15
Fungi N= 108, % OCCURRENCE = ..37%14
Herbs N= 108, % OCCURRENCE = 38%14
Insects, larvae N= 108, % OCCURRENCE = 26%10
Moss N= 108' % OCCURRENCE = 26%10
59
from till food type g/day kJ/day % of diet referencecomment
Bank vole Clethrionomys glareolus
Sep Nov Grass Hansson, 1971N= 108, % OCCURRENCE = 17%6
Berries N= 108, % OCCURRENCE = 11%4
Lignified stems N= 108, % OCCURRENCE = 5%2
Graminoids N= 108, % OCCURRENCE = 2%1
Lumbricids N= 108, % OCCURRENCE = 2%1
Dec Feb Seeds N= 46, % OCCURRENCE = 61%25
Insects, adults N= 46, % OCCURRENCE = 41%17
Herbs N= 46, % OCCURRENCE = 30%13
Moss N= 46, % OCCURRENCE = 26%11
Insects, larvae N= 46, % OCCURRENCE = 26%11
Grass N= 46, % OCCURRENCE = 17%7
Fungi N= 46, % OCCURRENCE = 17%7
Lignified stems N= 46, % OCCURRENCE = 13%5
Graninoids N= 46, % OCCURRENCE = 4%2
Molluscs N= 46, % OCCURRENCE = 2%1
Plants Hansson, 1985a% VOLUME in stomach contents from a centralEuropean lime-hornbeam forest, with few large seeds.Data from European beech forest similar, althoughseeds slightly greater (40%).
No seasonal data 50
Seeds 30
Small vertebrates 10
Insects, annelids& molluscs
5
Fungi 5
60
from till food type g/day kJ/day % of diet referencecomment
Bank vole Clethrionomys glareolus
Dead leaf matter Hansson, 1985bNo seasonal data 20
Seeds No seasonal data. Data from an English deciduousforest. Consumption of forb leaf and fungi similar toEuropean forests above. Little animal food eaten.
20
Dead leaf matter 20
Fruit 10
Vaccinium berries Hansson, 1985cNo seasonal data. Data from a conifer reforestation,which is representative of a conifer forest. Bark alsoappeared in the diet in appreciable but unquantifiedamounts.
No seasonal data 15
Chordate lichens 10
Fungi 10
Rosehips HBMFleshy fruits and seeds with a soft testa are eaten whenavailable. Leaves of woody plants preferred to those ofherbs. Juveniles eat less seed that adults. Only fleshof Rosehips is eaten, not skin.
No seasonal data
Jan Apr Dead leaves Watts, 1968% VOLUME in stomach contents40.5
Seeds mainly endosperm but traces of nettle and other seedtesta
26.5
Herbs eg primrose, blackberry bud19
May Jul Woody-plantleaves
epecially oak in May, privet in June and July, sycamoreJuly to September
46
Animal Not important in any other month15
61
from till food type g/day kJ/day % of diet referencecomment
Bank vole Clethrionomys glareolus
May Jul Herbs Watts, 196815
Seeds 11
Fungi 7
Sep Nov Seeds 27
Herbs 25
Fruits mainly blackberry, elderberry, hawthornberry21
Fungi 11
62
from till food type g/day kJ/day % of diet referencecomment
Brown hare Lepus europaeus
Oct Dec Wheat Frylestam, 1986aResults are expressed as % FREQUENCY in stomachcontents. Study of winter diet of hares (October -December) in relation to agricultural land use. Amongcultivated plants, wheat was most frequently eaten inmixed farmland and in monoculture land. Rape was wellrepresented in mixed farmland but was less ofteneaten. Rape was highly utilized in monoculture land.Wild sweet grasses play an important role. Agriculturalland use determines food intake. Sweden. Data splitinto monoculture (a), mixed farmland (b) and pastureland (c).
48.5
Rape 37.8
Deschampsiacaespitosa
4.7
Other herbs &woody plants
3
Bromus mollis 2.2
Graminae sp. 1.7
Deschampsiaflexuosa
1
Holcus lanatus 0.4
Hordeum vulgare 0.2
Poa annua 0.2
Lolium perenne 0.2
Oct Dec Wheat Frylestam, 1986bMixed farmland20.5
Graminae sp. 18
Rape 12.2
63
from till food type g/day kJ/day % of diet referencecomment
Brown hare Lepus europaeus
Oct Dec Bromus mollis Frylestam, 1986b12
Holcus lanatus 7.6
Lolium perenne 4.8
Deschampsiaflexuosa
4.2
Festuca rubra 2.6
Hordeum vulgare 2.6
Other herbs andwoody plants
2.4
Dactylis glomerata 2
Phleum pratense 1.8
Avena sativa 1.4
Poa annua 1.4
Poa pratensis 1.2
Agrostis tenuis 1
Deschampsiacaespitosa
1
Agropyron repens 0.8
Sinapis arvensis 0.5
Stellaria media 0.5
Other Poa sp. 0.3
Bellis perennis 0.2
Taraxacumvulgare
0.2
Vicia sp. 0.2
64
from till food type g/day kJ/day % of diet referencecomment
Brown hare Lepus europaeus
Oct Dec Poa trivialis Frylestam, 1986b0.2
Oct Dec Graminae sp. Frylestam, 1986cPasture21
Other herbs &woody plants
12
Festuca rubra 9.8
Deschampsiaflexuosa
9.7
Wheat 7.6
Lolium perenne 6.8
Dactylis glomerata 6
Deschampsiacaespitosa
5.2
Bromus mollis 3.4
Holcus lanatus 2.8
Poa pratensis 2.1
Agropyron repens 1.9
Phleum pratense 1.5
Other Poa sp. 1.4
Achillaeamillefolium
1.4
Poa annua 1.1
rape 1
Plantagolanceolata
0.9
65
from till food type g/day kJ/day % of diet referencecomment
Brown hare Lepus europaeus
Oct Dec Hieraciumpilosella
Frylestam, 1986c0.7
Agrostis tenuis 0.7
Poa trivialis 0.5
Carex sp. 0.4
Ranunculus sp. 0.3
Beta vulgaris 0.3
Trifolium repens 0.3
Experimental diet 500 3164 HBMUnder experimental conditions; diet not specified.No seasonal data
Grasses, herbs &arable crops
Prefers cereals during early stages of growth. Height ofcrop most important. Hares prefer shorter, more openvegetation.
66
from till food type g/day kJ/day % of diet referencecomment
Common dormouse Muscardinus avellanarius
May Leaves Richards et al., 1984% FREQUENCY in faecal pellets in S. Devon.Identifiable remains include insects (larval lepidoptera,aphids and other adult insects); green plant leaf,(honeysuckle, willow, hazel, parenchyma); pollen andspores, (honeysuckle, willowherb, spores); berries, nutsand bark.
45
Insects 20
May Pollen/spores Richards et al., 198435
Jun Insects 60
Leaves 30
Pollen/spores 5
Bark 5
Jul Leaves 60
Pollen/spores 20
Bark 10
Insests 10
Aug Nuts/berries 30
Leaves 30
Insects 20
Pollen/spores 10
Bark 10
Sep Nuts/berries 60
Leaves 30
Insects 5
67
from till food type g/day kJ/day % of diet referencecomment
Common dormouse Muscardinus avellanarius
Sep Pollen/spores Richards et al., 19845
Oct Nuts/berries 85
Leaves 10
Insects 5
68
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Natural diet 11.8 Aitchison, 1987Cites Tupikova (1949) stating that the energyrequirements of the common shrew in USSR at 15degrees C are 1.42 g food per g bodyweight per day.
No seasonal data
Natural diet Churchfield, 1982aMajor prey types in all seasons were adult coleopterans,insect larvae, araneids, opilionids and isopods. 41% ofprey taken were less than or equal to 5mm in bodylength. Data from analysis of faecal pellets. UK.
No seasonal data
Feb Feb Calliphora pupae 8.17 Churchfield, 1982b20 shrews, mean wt. 8.17g, maintained outdoors at 1-9degrees C. Animals consumed an average of 79.2%bodyweight.
May May Calliphora pupae 8.1 15 shrews, mean wt 8.1g maintained in the laboratory at20 degrees C. On average animals consumed 97% oftheir body weights.
Feb Lumbricids Churchfield, 1984% FAECES containing items. (Totals may exceed 100.)Study of shrews inhabiting watecress beds. UK.
62
Gastropods 54
Geophilomorphs 54
Tipulid larvae 46
Opilionids 39
Carabids 23
Coleopteranlarvae
15
Dipteran adults 15
Araneids 15
69
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Feb Acarines Churchfield, 19848
Lithobiomorphs 8
Plecopteran larvae 8
Coleopteran adults 8
Trichopteranlarvae
Cased larvae.8
Mar Lumbricids 72
Gastropods 67
Tipulid larvae 39
Araneids 28
Acarines 22
Staphylinids 17
Opilionids 17
Lepidopteranlarvae
17
Dipteran adults 17
Coleopteran adults 17
Formicids 11
Geophilomorphs 11
Plecopteran larvae 11
Isopods 6
Coleopteranlarvae
6
May Lumbricids 72
Dipteran adults 62
70
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
May Isopods Churchfield, 198456
Gastropods 50
Chrysomelids 28
Acarines 17
Coleopteran adults 17
Lepidopteranlarvae
11
Formicids 11
Carabids 11
Araneids 11
Geophilomorphs 11
Dermapterans 6
Hemipteran adults 6
Staphylinids 6
Jun Gastropods 78
Isopods 67
Dipteran adults 56
Lumbricids 50
Hemipteran adults 39
Formicids 28
Chrysomelids 28
Acarines 22
Coleopteran adults 17
71
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Jun Lepidopteranlarvae
Churchfield, 198411
Coleopteranlarvae
11
Araneids 11
Opilionids 11
Dipteran larvae 6
Lithobiomorphs 6
Jul Lumbricids 80
Dipteran adults 47
Acarines 47
Gastropods 47
Formicids 27
Dipteran larvae 27
Hemipteran adults 20
Carabids 20
Coleopteran adults 13
Araneids 13
Opilionids 13
Isopods 13
Geophilomorphs 7
Coleopteranlarvae
7
Sep Dipteran adults 95
Lumbricids 74
72
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Sep Gastropods Churchfield, 198453
Isopods 37
Coleopteran adults 21
Geophilomorphs 21
Araneids 21
Staphylinids 21
Formicids 16
Opilionids 16
Acarines 11
Chrysomelids 11
Dipteran larvae 5
Hemipteran adults 5
Asellus 5
Carabids 5
Oct Gastropods 78
Dipteran adults 33
Araneids 33
Lumbricids 33
Isopods 33
Geophilomorphs 22
Coleopteran adults 22
Dipteran larvae 22
Acarines 22
Dec Dipteran adults 90
73
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Dec Lumbricids Churchfield, 198440
Gastropods 30
Staphylinids 20
Araneids 10
Acarines 10
Geophilomorphs 10
Earthworms,gastropods &coleopterans
Churchfield, 1990No seasonal data 50
Invertebrates &neonatal mice
50.4 Hawkins & Jewell,1961
Captive animals, fed on a diet of maggots, mealworms,earthworms and neonatal mice required a daily foodintake of 1.80 - 2.30 kcal/g body weight.
No seasonal data
Invertebrates 58 HBM60 - 82% of prey eaten are >6 mm in length whichcorrelated with relative abundance of smallinvertebrates in soil and vegetation. Require 6.7 - 9.7kJ/g per day.
No seasonal data 60
Jan Dec Natural diet Pernetta, 1976aThe dominant components of the diet were earthworms,adult coleoptera and opiliones. Earthworms moreimportant in winter than in summer. Data from analysisof gut and stomach contents from shrews inhabitingOxfordshire grassland.
Jan Mar Lumbricidae Pernetta, 1976b% Wet weight in stomachs. Oxfordshire grassland25
74
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Jan Mar Coleoptera adults Pernetta, 1976b23
Opiliones 19
Aranae 13
Coleoptera larvae 6
Enchytraeidae earthworms4
Apr Jun Coleoptera adults 45
Lumbricidae 19
Coleoptera larvae 15
Opiliones 9
Aranae 7
Mollusca 4
Jul Sep Coleoptera adults 31
Lumbricidae 28
Opiliones 18
Coleoptera larvae 7
Aranae 6
Mollusca 2
Oct Dec Lumbricidae 47
Coleoptera adults 22
Enchytraeidae earthworms18
Aranae 4
Opiliones 3
Mollusca 2
75
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
May Sep Natural diet 58.1 Pernetta, 1976cSummer food requirement. As cited in Aitchison (1987)
Oct Apr Natural diet 61.1 Winter food requirement. As cited in Aitchison (1987)
Jan Feb Plant Rudge, 1968a% FREQUENCY in gut contents. Scotland.24
Coleoptera adults 16
Slugs 12
Diptera larvae 12
Hemiptera 8
Opiliones 8
Lumbricidae 4
Coleoptera larvae 4
Hymenoptera 4
Araneae 4
Acari 4
Apr May Plant 28
Coleoptera adults 17
Lumbricidae 14
Slugs 12
Hemiptera 8
Lepidoptera larvae 7
Diptera adults 5
Coleoptera larvae 3
Diptera larvae 2
76
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Apr May Other insectlarvae
Rudge, 1968a2
Araneae 1
Opiliones 1
Jun Aug Plants 22
Coleoptera adults 18
Lepidoptera larvae 11
Hemiptera 7
Diptera larvae 6
Araneae 6
Lumbricidae 6
Opiliones 5
Diptera adults 5
Other insectlarvae
4
Slugs 4
Thysanoptera 2
Acari 2
Hymenoptera 1
Sep Plant 27
Diptera adults 18
Opiliones 18
Coleoptera adults 18
Hymenoptera 9
77
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Sep Lumbricidae Rudge, 1968a9
Jan Plant Rudge, 1968b% FREQUENCY in gut contents. Wytham Woods, UK.15
Opiliones Oligolophus agrestis 9%, others 3%.12
Coleoptera Curculionidae 3%, others 9%.12
Coleoptera larvae 9
Diptera larvae Bibionidae 6%.6
Lumbricidae 6
Chilopoda Lithobiidae 6% .6
Discus rotundatus 6
Hemiptera 6
Diptera adults 6
Slugs 3
Other snails 3
Acari 3
Araneae 3
Other Insectalarvae
3
Hymenoptera 3
Mar Apr Plant 18
Other snails 13
Diptera larvae Bibionidae 3%, Tipulidae 5%, others 3%.11
Lumbricidae 10
Coleoptera Staphylinidae 10%.10
78
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Mar Apr Acari Rudge, 1968b10
Diptera adults 5
Araneae 5
Isopoda 3
Hymenoptera 3
Discus rotundatus 3
Chilopoda Geophilomorpha 3%.3
Slugs 3
Hemiptera 3
Other Insectalarvae
3
Coleoptera larvae 3
Jun Jul Plant 17
Opiliones Nemastoma lugubre 3%, others 9%.12
Coleoptera adults Staphylinidae 5%, Curculionidae 2%, others 5%.12
Slugs 11
Acari 8
Lumbricidae 8
Snails 6
Other Insectalarvae
6
Hemiptera 5
Chilopoda Geophilomorpha 3%, Lithobiidae 2%.5
Hymenoptera 3
79
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Jun Jul Diptera adults Rudge, 1968b2
Coleoptera larvae 2
Lepidoptera larvae 2
Other Insectaadults
2
Isopoda 2
Araneae 2
Sep Oct Plant 15
Opiliones Nemastoma lugubre 6%, others 7%.13
Lumbricidae 11
Hemiptera 9
Coleoptera Staphylinidae 1%, others 7%.8
Other snails 8
Diptera larvae Bibionidae 2%, others 4%.6
Other Insectalarvae
5
Discus rotundatus 5
Other Insectaadults
4
Acari 3
Hymenoptera 3
Slugs 3
Isopoda 1
Lepidoptera larvae 1
Chilopoda Lithobiidae 1%1
80
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Sep Oct Araneae Rudge, 1968b1
Mar May Lumbricidae Rudge, 1968c% FREQUENCY in 327 gut contents. Devon, UK.22
Plant 16
Diptera larvae 12
Chilopoda Geophilomorpha 4%, Lithobiidae 5%.9
Lepidoptera larvae 8
Coleoptera adults Curculionidae 3%, others 4%.7
Isopoda 5
Snails 4
Acari 4
Slugs 3
Siphonaptera 3
Coleoptera larvae 3
Araneae 3
Siphonaptera 3
Hymenoptera 1
Opiliones 1
Other larvae 1
Jun Aug Plant 17
Coleoptera adults Curculionidae 3%, others 11%.14
Lumbricidae 11
Snails 9
Chilopoda Geophilomorpha 1%, Lithobiidae 6%.7
81
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Jun Aug Isopoda Rudge, 1968c7
Opiliones 6
Acari 5
Araneae 5
Slugs 4
Lepidoptera larvae 3
Other larvae 3
Coleoptera larvae 3
Hymenoptera 2
Hemiptera 2
Siphonaptera 2
Diptera larvae 2
Diptera adults 1
Iulidae 1
Sep Nov Coleoptera adults Curculionidae 2%, others 16%.18
Plant 16
Lumbricidae 10
Hemiptera 7
Opiliones 7
Chilopoda Geophilomorpha 2%, Lithobiidae 5%.7
Araneae 6
Slugs 6
Other larvae 5
Isopoda 3
82
from till food type g/day kJ/day % of diet referencecomment
Common shrew Sorex araneus
Sep Nov Acari Rudge, 1968c3
Diptera adults 3
Snails 2
Diptera larvae 2
Hymenoptera 2
Iulidae 1
Siphonaptera 1
Coleoptera larvae 1
Dec Feb Plant 24
Slugs 16
Other larvae 13
Lumbricidae 13
Chilopoda Geophilomorpha 5%, Lithobiidae 3%.8
Opiliones 5
Isopoda 5
Diptera larvae 5
Coleoptera adults Not identified.5
Coleoptera larvae 3
Acari 3
83
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Sep Mar Acorns Caldwell et al, 1983% VOLUME in rumen contents. UK. Other foodsincluded: moss (0.1%), twigs (0.1%), bark (0.1%),bramble (0.3%), Juncus spp. (0.1%), herbs (0.2%),honeysuckle (0.1%), horse chestnut (0.2%) and wildrose (0.1%).
49
Grasses 40
Corn & chaff 4
Broadleaved treeleaves, dead
3.4
Crab apple 1.7
Broadleavedtrees, live
0.7
Grasses HBMHerbs and broad leaf browse also make significantcontribution.
No seasonal data 60
Jan Graminids Jackson, 1977% VOLUME in rumen contents collected during a 3year study. New Forest, UK. Throughout the year, 43%of the diet consisted of graminids, ranging from 21% inwinter to 67% in spring/summer.
21
Bramble & rose 17
Calluna 16
Conifers 14
Holly 12
Ivy 8
Bilberry 2
Ferns 2
84
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Jan Acorn Jackson, 19772
Fungi 1
Bryophytes 1
Dead broadleaves 1
Herbs 1
Feb Graminids 25
Calluna 24
Holly 17
Conifers 14
Bramble & rose 7
Bramble & rose 7
Ivy 4
Bilberry 2
Bryophytes 2
Bark 1
Ferns 1
Other material 1
Fungi 1
Herbs 1
Dead broadleaves 1
Mar Graminids 59
Calluna 16
Holly 9
Conifers 7
85
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Mar Bilberry Jackson, 19774
Bramble & rose 2
Bramble & rose 2
Herbs 1
Ivy 1
Bryophytes 1
Apr Graminids 67
Holly 7
Herbs 6
Broadleaf trees 4
Calluna 4
Bilberry 4
Calluna 3
Bramble & rose 3
Bark 2
Ivy 1
Bryophytes 1
Dead broadleaves 1
Other dwarfshrubs
1
Conifers 1
Ferns 1
May Jul Graminids 63
Broadleaf trees 14
86
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
May Jul Herbs Jackson, 19776
Ivy 4
Holly 4
Calluna 4
Bramble & rose 3
Bilberry 2
Bryophytes 1
Aug Graminids 57
Herbs 12
Bramble & rose 12
Broadleaf trees 11
Calluna 3
Holly 3
Bilberry 1
Ivy 1
Sep Graminids 58
Acorn 13
Bramble & rose 10
Bramble & rose 7
Herbs 7
Broadleaf trees 5
Ivy 2
Calluna 2
Dead broadleaves 1
87
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Sep Fungi Jackson, 19771
Crab apple 1
Other fruits & nuts 1
Holly 1
Oct Graminids 33
Acorn 32
Bramble & rose 10
Mast 9
Fungi 6
Broadleaf trees 3
Herbs 2
Dead broadleaves 1
Calluna 1
Gorse 1
Nov Graminids 26
Acorn 19
Bramble & rose 10
Conifers 8
Calluna 8
Dead broadleaves 8
Bilberry 5
Ivy 4
Broadleaf trees 4
Herbs 2
88
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Nov Mast Jackson, 19772
Holly 2
Ferns 1
Fungi 1
Bryophytes 1
Crab apple 1
Dec Graminids 21
Conifers 17
Calluna 16
Acorn 14
Bramble & rose 8
Holly 7
Ivy 4
Dead broadleaves 3
Bilberry 2
Fungi 2
Ferns 1
Mast 1
Broadleaf trees 1
Herbs 1
Feb Mosses Putman & Hemmings,1986
% VOLUME of fragments in faeces corrected fordigestibility. New Forest, UK.
44.5
Other grasses 26.2
89
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Feb Ulex Putman & Hemmings,1986
12.4
Bracken & ferns 3
Conifers 3
Other shrubs 2.9
Agrostis setacea 1.8
Calluna/Erica 1.5
Acorn 1.5
Agrostis tenuis 1.3
Forbs 1.2
Juncus sp. 0.6
Apr Other grasses 33.6
Conifers 21
Bracken & ferns 17.7
Mosses 11
Ulex 9.7
Juncus sp. 2.2
Other shrubs 1.2
Forbs 1.2
Agrostis setacea 1.1
Quercus 0.7
Agrostis tenuis 0.6
Jun Other grasses 37
Agrostis tenuis 23
90
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Jun Agrostis setacea Putman & Hemmings,1986
13
Bracken & ferns 12
Forbs 3.5
Juncus sp. 3.4
Ulex 2.3
Conifers 2.1
Mosses 1.6
Quercus 0.8
Calluna/Erica 0.6
Other shrubs 0.6
Aug Other grasses 38
Agrostis setacea 14
Agrostis tenuis 9.9
Bracken & ferns 8
Forbs 7.7
Other shrubs 5.7
Quercus 5.2
Juncus sp. 3.9
Mosses 3.6
Conifers 3
Ulex 0.9
Oct Mosses 42
Querus 17
91
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Oct Other grasses Putman & Hemmings,1986
16
Other shrubs 8.5
Agrostis tenuis 3.1
Ulex 2.9
Bracken & ferns 2.7
Agrostis setacea 2.7
Forbs 1.9
Betula 1.9
Juncus sp. 0.7
Dec Mosses 32
Ulex 19.6
Conifers 19
Other shrubs 12
Other grasses 5.5
Calluna/Erica 3.3
Acorn 2.7
Quercus 1.6
Hedera 1.4
Agrostis setacea 1.1
Rubus 0.9
Agrostis tenuis 0.6
92
from till food type g/day kJ/day % of diet referencecomment
Fallow deer Dama dama
Grasses Putman et al., 1993Fallow deer in New Forest shown to be predominantlygrazers, grasses accounting for approx. 70% of annualforage intake, with remainder of diet being sedges,rushes, heather and broadleaved browse.
No seasonal data 70
93
from till food type g/day kJ/day % of diet referencecomment
Feral cattle Bos taurus
Jan Grasses Putman, 1986% FREQUENCY of plant cuticular fragments identifiedin faeces. New Forest, UK.
75
Heather 21
Other 3
Broadleaves 1
Feb Grasses 67
Heather 27
Other 5
Broadleaves 1
Mar Grasses 71
Heather 24
Other 4
Broadleaves 1
Apr Grasses 80
Other 10
Heather 9
Broadleaves 1
May Grasses 80
Heather 14
Other 5
Herbs 1
Jun Grasses 83
Heather 12
Other 5
94
from till food type g/day kJ/day % of diet referencecomment
Feral cattle Bos taurus
Jul Grasses Putman, 198681
Heather 13
Other 5
Molinia 1
Aug Grasses 70
Heather 18
Other 11
Broadleaves 1
Sep Grasses 65
Heather 23
Other 8
Broadleaves 2
Herbs 1
Molinia 1
Oct Grasses 69
Heather 19
Other 9
Broadleaves 1
Herbs 1
Molinia 1
Nov Grasses 69
Heather 21
Other 7
Broadleaves 2
95
from till food type g/day kJ/day % of diet referencecomment
Feral cattle Bos taurus
Nov Molinia Putman, 19861
Dec Grasses 65
Heather 22
Other 10
Broadleaves 1
Herbs 1
Molinia 1
Calamagrostis Williams et al, 1973% VOLUME of plant fragments in faeces of 10 Gallowaysteers grazed on Woodwalton fen. Calamagrostis wasthe most consistent contributor to diet, ranging from 18 -47%. Other species include Festuca and Juncus inearly winter, Phragmites and Poa pratensis in mid-summer.
No seasonal data 33
Calamagrostis Williams et al, 1974Data obtained by faecal analysis of 10 Galloway steersgrazed on Woodwalton fen. Calamagrostis was themost consistent contributor to diet, ranging from 18 -47%. Other species include Festuca and Juncus inearly winter, Phragmites and Poa pratensis in mid-summer.
No seasonal data 33
96
from till food type g/day kJ/day % of diet referencecomment
Feral goat Capra hircus
Bullock, 1985aFaecal analysis, Scotland, UK a) Cairnsmore, b)Langholm hills, c) Nether Hindhope.At Cairnsmore, woody shrubs, especially Calluna, themain dietary component. Ericaceous shrubs and Myricaeaten mainly late autumn, winter and early spring. FromMay to September diet dominated by sedges(Trichophorum) and grasses (Molinia). BetweenOctober and April Festuca and Nardus were the mostfrequently recorded grasses.
No seasonal data
Bullock, 1985bLangholm Hills, during summer, goats ate more sedges(Trichophorum and Carex) and less grass than didsheep and also showed a spring peak, in the use ofrushes (Juncus). In autumn and winter months goatdiets contained more ericaceous shrubs than did sheepdiets. Consumption of ferns (Pteridium) was generallylow but reached a peak of 24% in October.
No seasonal data
Bullock, 1985cNether Hindhope conifer plantation. Conifer needlespredominated in February. Summer diet consistedmainly of rushes (Juncus) and grasses (Festuca andMolinia).
No seasonal data
HBMSummer diet predominated by grasses, sedges andrushes. Increased use of dwarf shrubs, gorse and otherbrowse in winter. May eat seaweed and bark-stripwillow, rowan, oak, alder and conifers.
No seasonal data
97
from till food type g/day kJ/day % of diet referencecomment
Feral sheep Ovis aries
Bullock, 1985aFaecal analysis, Scotland, UK a) Cairnsmore, b)Langholm hills, c) Nether Hindhope.At Cairnsmore, woody shrubs, especially Calluna, themain dietary component. Ericaceous shrubs and eatenmainly late autumn, winter and early spring. From Mayto September diet dominated by sedges (Trichophorum)and grasses (Molinia). Between October and AprilFestuca and Nardus were the most frequently recordedgrasses.
No seasonal data
Bullock, 1985bNether Hindhope. Grasses were the most importantfood item for sheep the year round. Festucapredominated although Molinia reached 17% in thesummer.
No seasonal data
Langholm hills. Grasses dominated sheep diets fromMay to October. Between June ans August, Molinia wasmain species recorded, wheile Festuca predominated inMay, September and October.
HBMGrasses predominate in summer with an increasingproportion of heather towards and through the winter forSoay sheep. Boreray sheep graze all year.
No seasonal data
98
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Jan Dec Brachypodiumsylvaticum
Evans, 1973aData from gut analysis. No % figures given. On a 10yrold tree plantation, the main grasses taken were,Festuca rubra, Arrhenatherum elatius, Brachypodiumpinnatum, Zena erecta and Festuca rubra. UK.
Oct May Brachypodiumsylvaticum
Evans, 1973bData from gut analysis. No % figures given. On an areaof ungrazed limestone grassland, the principal grassspecies taken were Dactylis glomerata, Holcus lanatus,Arrhenatherum elatius, Poa trivialis, Poa pratensis,Brachypodium sylvaticum and Festuca rubra. UK.
Mar Holcus lanatus Faber & Ma, 1986a% FRESH WEIGHT in stomach contents. Budel,Netherlands.
87
Agrostis capillaris 6.6
Undetermineddicotyledons
2.5
Elymus repens Couch grass1.6
Undeterminedplant material
1
Poa angustifoliaand P. pratensis
0.9
Fungi 0.3
Jun Holcus lanatus 37
Undetermineddicotyledons
18
Lychnis flos-cuculi 17.8
Agrostis capillaris 8.4
99
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Jun Poa angustifolia &P. pratensis
Faber & Ma, 1986a6.7
Cytisus scoparius(flower)
Broom4.7
Undeterminedplant material
3
Animal material 2
Aug Poa angustifolia &P. pratensis
33
Undetermineddicotyledons
29
Holcus lanatus 12
Undeterminedplant material
8.3
Elymus repens Couch grass7.4
Agrostis capillaris 5
Seeds 4.9
Animal material 1.2
Oct Holcus lanatus 33
Elymus repens Couch grass19
Undetermineddicotyledons
17
Poa angustifolia &P. pratensis
14
Agrostis capillaris 13
100
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Oct Undeterminedplant material
Faber & Ma, 1986a2
Seeds 0.7
Other 0.3
Dec Holcus lanatus 63
Agrostis capillaris 33
Undeterminedplant material
2.5
Other 0.7
Undetermineddicotyledons
0.3
Mar Deschampsiaflexuosa
Faber & Ma, 1986b% FRESH WEIGHT in stomach contents. Arnhem,Netherlands.
Wavy hair grass
92
Hypnumcypressiforme
2.9
Undeterminedplant material
2.2
Undetermineddicotyledons
1
Vaccinium myrtilla 0.8
Lignified stems 0.7
Jun Deschampsiaflexuosa
Wavy hair grass77
101
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Jun Hypnumcypressiforme
Faber & Ma, 1986b7.3
Vaccinium myrtilla 6
Undeterminedplant material
4.4
Lignified stems 3.4
Undetermineddicotyledons
0.8
Animal 0.4
Aug Deschampsiaflexuosa
Wavy hair grass72
Hypnumcypressiforme
13
Vacciniummyrtillus berry
4.3
Lignified stems 4.2
Undeterminedplant material
3
Carex arenaria 1.2
Animal 1.1
Undetermineddicotyledons
1
Vaccinium myrtilla 1
Oct Deschampsiaflexuosa
Wavy hair grass78
Undeterminedplant material
11
102
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Oct Undeterminedseed
Faber & Ma, 1986b3
Undetermineddicotyledons
2.4
Hypnumcypressiforme
2.3
Vaccinium myrtilla 1.7
Lignified stems 1
Fungi 0.1
Dec Deschampsiaflexuosa
100
63.7 Ferns, 1976aLaboratory study of captive animals.No seasonal data
Apr Mar Unidentifiedpanicoid grasses
Ferns, 1976b% VOLUME in faeces. Mean over 12 months.Dicotyledons appeared in small quantities throughoutthe year. UK.
28
Festuca rubra Red fescue20
Mosses 16
Other 9
Anthoxanthumodoratum
Sweet vernal grass6.8
Agrostis canina 4.3
Juncus spp. 4.2
Cynosuruscristatus
3.8
103
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Apr Mar Poa trivialis Ferns, 1976b3.7
Dactylis glomerata 1.9
Holcus lanatus 1.2
Phleum pratense 1
Mar Mar Grass Hansson, 1971% NUMBER of items found in coecae of field volescaught in Southern Swedish forest.N=21 %OCCURRENCE=100%
33
Lignified stems N=21 %OCCURRENCE=43%14
Graminoids N=21 %OCCURRENCE=33%11
Herbs N=21 %OCCURRENCE=24%8
Insects, larvae N=21 %OCCURRENCE=19%6
Moss N=21 %OCCURRENCE=10%3
Seeds N=21 %OCCURRENCE=5%2
Roots N=21 %OCCURRENCE=5%2
Apr May Grass N=69 %OCCURRENCE=99%34
Moss N=69 %OCCURRENCE=51%18
Herbs N=69 %OCCURRENCE=38%13
Insects, larvae N=69 %OCCURRENCE=20%7
Lignified stems N=69 %OCCURRENCE=16%6
Graminoids N=69 %OCCURRENCE=26%4
Insects, adults N=69 %OCCURRENCE=6%2
Roots N=69 %OCCURRENCE=1%1
Seeds N=69 %OCCURRENCE=1%1
104
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Jun Aug Grass Hansson, 1971N=80 %OCCURRENCE=95%33
Herbs N=80 %OCCURRENCE=80%26
Moss N=80 %OCCURRENCE=38%13
Graminoids N=80 %OCCURRENCE=33%11
Insects, larvae N=80 %OCCURRENCE=14%5
Seeds N=80 %OCCURRENCE=11%4
Insects, adults N=80 %OCCURRENCE=13%4
Fungi N=80 %OCCURRENCE=8%3
Lignified stems N=80 %OCCURRENCE=4%1
Sep Nov Grass N=193 %OCCURRENCE=100%44
Herbs N=193 %OCCURRENCE=36%16
Moss N=193 %OCCURRENCE=24%11
Graminoids N=193 %OCCURRENCE=17%7
Seeds N=193 %OCCURRENCE=16%7
Lignified stems N=193 %OCCURRENCE=9%4
Insects, larvae N=193 %OCCURRENCE=9%4
Fungi N=193 %OCCURRENCE=9%4
Insects, adults N=193 %OCCURRENCE=6%3
Roots N=193 %OCCURRENCE=2%1
Dec Feb Grass N=55 %OCCURRENCE=98%37
Graminoids N=55 %OCCURRENCE=60%23
Herbs N=55 %OCCURRENCE=38%15
Lignified stems N=55 %OCCURRENCE=18%7
Moss N=55 %OCCURRENCE=18%7
105
from till food type g/day kJ/day % of diet referencecomment
Field vole Microtus agrestis
Dec Feb Seeds Hansson, 1971N=55 %OCCURRENCE=9%3
Insects, adults N=55 %OCCURRENCE=5%2
Insects, larvae N=55 %OCCURRENCE=2%1
Monocotyledons Rogers, 1990% VOLUME in gut contents from a population living inoilseed rape fields. Plant material made up more than95% of the diet. UK.
No seasonal data 82
Oilseed rape 3
106
from till food type g/day kJ/day % of diet referencecomment
Fox Vulpes vulpes
Small mammals Borkowski, 1994% FREQUENCY of prey items in faeces from analysisof 144 scats. Pine voles were the most important smallmammal. Poland.
No seasonal data 54
Medium mammals Marmot was the most frequently eaten medium-sizedmammal.
43
Mar May Water voles Ferrari & Weber, 1995% FREQUENCY of prey items in faeces. Switzerland.38
Grass 25
Exploitable scraps Domestic food scraps and cow placentae.9.9
Others 9.1
Earthworms 7.4
Wild fruits 5
Domestic stock Hens, geese, turkeys and domestic rabbits.3.3
Small mammals Apodemus sp., Clethrionomys glareolus, Sorex minutusand Mircotus arvalis.
2.5
Jun Aug Grass Possibly ingested in the course of feeding onmammalian and invertebrate prey.
32
Water voles 23
Others 14
Wild fruits 8.9
Small mammals 7.1
Domestic stock 5.4
Exploitable scraps 5.4
Earthworms 3.6
Sep Nov Wild fruits 31
107
from till food type g/day kJ/day % of diet referencecomment
Fox Vulpes vulpes
Sep Nov Earthworms Ferrari & Weber, 199524
Grass 12
Others 10
Water voles 7
Exploitable scraps 6.9
Small mammals 6.9
Domestic stock 2.3
Dec Feb Grass 37
Water voles 23
Earthworms 15
Exploitable scraps 10
Wild fruits 8.2
Small mammals 4
Domestic stock 2.5
Others 1.3
Lagomorpha Goszczynski, 1986% WEIGHT in faeces. 7-year study. Poland. Smallmammals and fruits eaten mainly in summer andautumn. Also eaten: frogs (0.03%) and small birds(0.09%). Rabbits and hares taken all year but peak inspring.
No seasonal data 26
Microtus arvalis 22
Gallus domesticus 14
Phasianuscolchicus
7.3
108
from till food type g/day kJ/day % of diet referencecomment
Fox Vulpes vulpes
Fruits Goszczynski, 1986No seasonal data 6.5
Fruits 6.5
Clethrionomysglareolus
4.2
Columba sp. 2.7
Apodemus sp. 2
Microtusoeconomus
2
Sus scrofa(carrion)
1.91
Capreoluscapreolus (carrion)
1.2
Canis familiaris(carrion)
0.91
Pitymyssubterraneus
0.9
Talpa europea &Sorex sp.
0.87
Perdix perdix 0.7
Insects Spring.0.6
Felis catus(carrion)
0.5
Sciurus vulgaris 0.12
109
from till food type g/day kJ/day % of diet referencecomment
Fox Vulpes vulpes
Scavenged meat,bones & fat
Harris, 1981ai% VOLUME in stomach contents. Non-food items(mostly lawn grasses) made up 25% of stomachcontents. Suburban study over 6 years. London, UK.Data separated into adults (a), culled over the wholeyear but mainly October - June; and cubs (c), culledApril - September. Data is further separated to take intoaccount differences in diet for foxes in the inner zone (i)(>5km within Greater London Council boundary; andthose in the outer zone (o) (<5km within GLC boundary).
No seasonal data 27
Jan Dec Wild birds 15
Other scavengeditems
13
Wild mammals 10.1
Insects 10.1
Earthworms 9.7
Fruit & vegetables 8.5
Pet birds 5.1
Pet mammals 1.7
Jan Dec Scavenged meat,bones & fat
Harris, 1981ao22
Wild mammals 16
Earthworms 14
Wild birds 14
Other scavengeditems
9.2
Insects 8.7
110
from till food type g/day kJ/day % of diet referencecomment
Fox Vulpes vulpes
Jan Dec Fruit & vegetables Harris, 1981ao6.8
Pet birds 6.3
Pet mammals 3.8
Apr Sep Wild birds Harris, 1981ci27
Insects 14
Scavenged meat,bones & fat
14
Wild mammals 10
Earthworms 10
Fruit & vegetables 8.1
Other scavengeditems
7.3
Pet mammals 4.7
Pet birds 4.3
Apr Sep Wild birds Harris, 1981co28
Insects 18
Earthworms 13
Scavengedbones, meat & fat
13
Wild mammals 10
Fruit & vegetables 8.4
Other scavengeditems
4.7
111
from till food type g/day kJ/day % of diet referencecomment
Fox Vulpes vulpes
Apr Sep Pet mammals Harris, 1981co3.2
Pet birds 1.8
Experimental diet 3448 HBMCaptive foxes have been found to require 507 kJ per kgbody wt.
No seasonal data
Sep Aug Small mammals Reynolds & AebischerApodemus 2.2%, Microtus 5.3%10.8
Sep Aug Lagomorph Reynolds &Aebischer, 1991
% FRESH WEIGHT extrapolated from faecal analysis.Rural, UK.rabbit 21%, hare 10%, Unidentified 11%
42
Birds gamebirds 17%, passerines 3%, pigeons 4%,unidentified 5%
30
Miscellaneousvertebrate
deer 1.6%, sheep 5%.14
Beetles 2.5
Earthworms 2
Brown hares Reynolds & Tapper,1995a
Study in mixed farming area of S. England. The numberof brown hares eaten by foxes accounted for 76 - 100%of the annual production. The hare population could notwithstand more than a very low addititonal mortalitywithout declining.
No seasonal data
112
from till food type g/day kJ/day % of diet referencecomment
Fox Vulpes vulpes
Rabbit Reynolds & Tapper,1995b
% VOLUME in scats. Mixed farming area of S England.Two thirds of fox diet consisted of vertebrate prey in therange 0.3 - 3.0 kg. Small rodents, fruit and vegetableswhich are V. important in other habitats are <10% eachhere because of presence of abundant game species;pheasant, brown hare, grey partridge and redleggedpartridge.
No seasonal data 37
Galliformes Mainly game birds16.3
Small mammals 11.2
Hare 11.1
Large mammals 9.4
Passeriformes 4.5
Colombiformes 4
Beetles 2.2
Fruit 2.1
Earthworms 2
Birds eggs 0.2
113
from till food type g/day kJ/day % of diet referencecomment
Grey squirrel Sciurus carolinensis
177 Bolls & Perfect, 1972Data obtained by respirometry. (From Knee, 1983).Laboratory study.
No seasonal data
Seed 70 HBM60 - 80 g seed eaten daily. Laboratory conditions maynot be truly representative of wild animals.
No seasonal data
175 Innes, 1978Data obtained by direct daily calorimetry. USA. (FromKnee, 1983). Laboratory study.
No seasonal data
Jan 400 Knee, 1983Seasonal daily assimilation of energy of captive greysquirrels.
Feb 310
Apr 350
May 460
Jun 520
Jul 480
Sep 700
Oct 640
Nov 620
Dec 510
Mar May Flowers, buds,coarse plant foods
MacKinnon, 1976Analysis method not given. (From Moller, 1983). UK.57
Animal matter 7
May Insects 22
114
from till food type g/day kJ/day % of diet referencecomment
Grey squirrel Sciurus carolinensis
Jun Beech vasculartissue
MacKinnon, 197682
Aug Deciduous seed& fruit
45
Flowers, buds,coarse plant food
Acer sp. and Quercus sp. particularly favoured flowers.Shoots of hazel, hawthorn. sycamore, beech and horsechestnut eaten. Bark stripped to reach cambial tissue,particularly from beech, Acer sp., oak, birch and ash.
40
Jul Insects 10
Aug Animal matter 9
Fungi 1
Sep Nov Deciduous seed& fruit
98
Fungi 1
Dec Feb Deciduous seed& fruit
96
Flowers, buds,coarse plant foods
3
Mar May Deciduous seed& fruit
MacKinnon. 197631
Fungi 1
416 Short & Duke, 1971Data obtained by daily assimilation gravimetry. (FromKnee, 1983). Laboratory study.
No seasonal data
115
from till food type g/day kJ/day % of diet referencecomment
Grey squirrel Sciurus carolinensis
Deciduous seed Taylor, 1969Some deciduous seeds eaten before the seed is matureeg. beechnuts taken in mid-summer, hazelnuts July andAugust. (From Moller, 1983). UK.
No seasonal data
116
from till food type g/day kJ/day % of diet referencecomment
Harvest mouse Micromys minutus
Sep Jan Fruit Dickman, 1986a% VOLUME in faeces. Urban populations from 3different habitats: (a) Orchard; (b) Long grass; (c)Scrub. UK. Study from September - January.
27
Seed 20.2
Monocotyledonleaf
12.4
Insect 12
Fungus 9.7
Dicotyledon leaf 9.4
Other Includes fibrous material, plant hooks and hairs.4.4
Other invertebrate 3.8
Vertebrate 0.3
Moss 0.2
Root 0.2
Sep Jan Seed Dickman, 1986bLong grass habitat.45
Monocotyledonleaf
21
Insect 16
Fruit 6.3
Other 4.3
Dicotyledon leaf 3.8
Fungus 3.3
Root 1
Other invertebrate 0.3
117
from till food type g/day kJ/day % of diet referencecomment
Harvest mouse Micromys minutus
Sep Jan Seed Dickman, 1986cScrub habitat.30
Dicotyledon leaf 19
Insect 16
Fruit 15
Monocotyledonleaf
6.8
Root 4.2
Moss 3.8
Fungus 2.6
Other 2.4
Vertebrate 0.2
Captive diet 36.12 Gorecki, 1971Average daily metabolic rate of captive mice: 1.07kcal/g/day.
No seasonal data
Nov Dec Oats, maize,sunflower seed &mealworms
2.1 Hawkins & Jewell,1962
Data given is wet weight from captive communallyhoused animals.
Oats, maize,sunflower seed &mealworms
1.4 28.5 Data given is dry weight. Equivalent to 0.97 kcal/g/day.
Captive diet 2 30 HBM29.3 - 36 kJ required per day, confirmed by bothrespiratory and dietary studies.
No seasonal data
118
from till food type g/day kJ/day % of diet referencecomment
Hedgehog Erinacus europaeus
Natural diet 70 HBMAverage nightly intake.No seasonal data
Lepidoptera -larvae
Yalden, 1976% WEIGHT in 137 stomachs. East Anglia, UK.No seasonal data 26
Scarabaeoidea 17
Lumbricidae 13
Aves (eggs) 11
Carabidae 8
Mammalia 5.3
Mollusca - slugs 4.1
Diplopoda 3.4
Dermaptera 3.3
Apoidea 2.7
Other Coleoptera 2.4
Tipulidae (larvae) 1.2
Aves (feathers) 1.1
Mollusca - snails 0.6
Other items 0.4
OtherHymenoptera
0.3
Araneida 0.1
Opiliones 0.1
Isopoda 0.1
Chilopoda 0.1
119
from till food type g/day kJ/day % of diet referencecomment
Hedgehog Erinacus europaeus
Diptera Imagines Yalden, 1976No seasonal data 0.1
Apr Lepidoptera(larvae)
Yalden, 1976a% STOMACHS containing item. (Totals for any onemonth may exceed 100). UK. Figures for Mammaliaand eggs likely to be inflated since traps are usuallybaited with rabbits or addled eggs.
64
Carabidae 58
Diplopda 46
Other Coleoptera 41
Dermaptera 35
Lumbricidae 35
Aves (egg) 12
Mollusca - slugs 12
Aves (feather) 12
Scarabeoidae 12
Apoidea 6
Mammalia 6
May Dermaptera 70
Carabidae 65
Lepidoptera(larvae)
59
Other Coleoptera 43
Diplopda 41
Lumbricidae 37
Mollusca - slugs 26
120
from till food type g/day kJ/day % of diet referencecomment
Hedgehog Erinacus europaeus
May Apoidea Yalden, 1976a17
Aves (egg) 15
Mammalia 13
Tipulidae (larvae) 11
Scarabeoidae 11
Aves (feather) 11
Jun Carabidae 86
Dermaptera 66
Aves (feather) 46
Lumbricidae 40
Diplopda 33
Other Coleoptera 26
Mollusca - slugs 20
Scarabeoidae 13
Aves (egg) 7
Tipulidae (larvae) 7
Lepidoptera(larvae)
7
Jul Carabidae 63
Diplopda 56
Scarabeoidae 42
Apoidea 35
Other Coleoptera 35
Aves (feather) 28
121
from till food type g/day kJ/day % of diet referencecomment
Hedgehog Erinacus europaeus
Jul Lumbricidae Yalden, 1976a21
Aves (egg) 21
Dermaptera 21
Mammalia 21
Lepidoptera(larvae)
14
Mollusca - slugs 14
Aug Lepidoptera(larvae)
71
Carabidae 47
Scarabeoidae 47
Lumbricidae 43
Dermaptera 43
Diplopda 29
Mollusca - slugs 29
Aves (egg) 14
Aves (feather) 14
Other Coleoptera 14
Apoidea 14
Sep Carabidae 88
Lepidoptera(larvae)
75
Dermaptera 63
Mollusca - slugs 50
Lumbricidae 37
122
from till food type g/day kJ/day % of diet referencecomment
Hedgehog Erinacus europaeus
Sep Diplopda Yalden, 1976a37
Mammalia 25
Other Coleoptera 25
Apoidea 12
Scarabeoidae 12
Aves (feather) 12
Oct Dermaptera 71
Lepidoptera(larvae)
62
Diplopda 48
Lumbricidae 43
Carabidae 33
Scarabeoidae 33
Other Coleoptera 24
Mollusca - slugs 19
Mammalia 5
Apoidea 5
Aves (feather) 5
Nov Other Coleoptera 33
Mollusca - slugs 33
Lumbricidae 33
Dermaptera 33
Lepidoptera(larvae)
33
123
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Mar May Grasses HBM% VOLUME of cuticular fragments in faeces. NewForest ponies.
48
Broad-leavedbrowse
21
Gorse 7
Heather 5
Jun Aug Grasses 90
Heather 1
Broad-leavedbrowse
1
Sep Nov Grasses 83
Heather 2
Broad-leavedbrowse
1
Dec Feb Grasses 56
Broad-leavedbrowse
15
Gorse 12
Heather 7
Jan Other Gramineae Putman et al, 1987% VOLUME of plant cuticular fragments identified infaeces. New Forest ponies. UK.
29
Tree leaves 19
Agrostis curtisii 15
Ulex sp. 12
124
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Jan Mosses Putman et al, 19878.5
Calluna vulgaris 6.7
Molinia caerulea 2.5
Agrostis capillaris 2
Festuca rubra 1.3
Pteridiumaquilinum
1.2
Agrostis canina 1.2
Juncus sp. 0.8
Carex sp. 0.7
Eriophorum sp. 0.5
Other 0.3
Feb Tree leaves 26
Other Gramineae 22
Ulex sp. 13
Agrostis curtisii 13
Mosses 13
Calluna vulgaris 7.3
Pteridiumapuilinum
2
Carex sp. 1
Molinia caerulea 1
Agrostis capillaris 0.7
Juncus sp. 0.5
125
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Feb Agrostis canina Putman et al, 19870.3
Eriophorum sp. 0.3
Festuca rubra 0.2
Other 0.2
Mar Other Gramineae 25
Tree leaves 25
Mosses 14
Agrostis curtisii 11
Ulex sp. 10.2
Agrostis canina 6.2
Calluna vulgaris 5.2
Juncus sp. 1.2
Pteridiumapuilinum
0.8
Carex sp. 0.7
Agrostis capillaris 0.5
Other 0.5
Molinia caerulea 0.3
Eriophorum sp. 0.3
Festuca rubra 0.2
Apr Other Gramineae 37
Agrostis curtisii 21
Mosses 15
Tree leaves 13
126
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Apr Agrostis canina Putman et al, 19874.8
Calluna vulgaris 3.2
Juncus sp. 1.3
Ulex sp. 1.2
Agrostis capillaris 1
Carex sp. 0.8
Other 0.5
Festuca rubra 0.5
Pteridiumapuilinum
0.3
Molinia caerulea 0.3
May Other Gramineae 54
Molinia caerulea 17
Agrostis curtisii 7.5
Agrostis capillaris 6.7
Festuca rubra 3.8
Mosses 3
Tree leaves 1.7
Eriophorum sp. 1.4
Agrostis canina 1.2
Other 1.2
Carex sp. 1
Calluna vulgaris 1
Juncus sp. 0.7
127
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Jun Other Gramineae Putman et al, 198748
Molinia caerulea 22
Agrostis capillaris 10.3
Agrostis curtisii 6.7
Mosses 4
Pteridiumapuilinum
1.7
Festuca rubra 1.3
Carex sp. 1.3
Agrostis canina 1.2
Tree leaves 1
Juncus sp. 0.8
Eriophorum sp. 0.7
Calluna vulgaris 0.5
Forbs 0.2
Jul Other Gramineae 50
Molinia caerulea 24
Agrostis capillaris 12
Agrostis curtisii 4.8
Pteridiumapuilinum
2.2
Eriophorum sp. 1.3
Other 1.2
Juncus sp. 0.8
128
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Jul Calluna vulgaris Putman et al, 19870.7
Mosses 0.7
Carex sp. 0.5
Agrostis canina 0.5
Festuca rubra 0.5
Tree leaves 0.2
Forbs 0.2
Aug Other Gramineae 50
Molinia caerulea 17
Agrostis curtisii 11
Agrostis capillaris 7.8
Pteridiumapuilinum
6.5
Mosses 1.5
Other 1.5
Agrostis canina 1
Juncus sp. 1
Tree leaves 1
Calluna vulgaris 0.8
Festuca rubra 0.7
Carex sp. 0.7
Eriophorum sp. 0.5
Forbs 0.3
Sep Other Gramineae 45
129
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Sep Agrostis curtisii Putman et al, 198724
Pteridiumapuilinum
9.7
Molinia caerulea 7
Agrostis capillaris 3.3
Agrostis canina 2.5
Mosses 2.2
Calluna vulgaris 1.7
Tree leaves 1.3
Festuca rubra 1
Juncus sp. 0.7
Eriophorum sp. 0.7
Other 0.7
Ulex sp. 0.5
Carex sp. 0.2
Oct Other Gramineae 39
Agrostis curtisii 32
Pteridiumapuilinum
6.5
Mosses 4.5
Tree leaves 3.3
Molinia caerulea 2.7
Agrostis canina 2.7
Agrostis capillaris 2.5
130
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Oct Calluna vulgaris Putman et al, 19872.5
Eriophorum sp. 1.3
Ulex sp. 1.2
Other 0.7
Carex sp. 0.7
Festuca rubra 0.5
Juncus sp. 0.2
Nov Agrostis curtisii 33
Other Gramineae 30
Tree leaves 11
Mosses 7.2
Calluna vulgaris 5.3
Ulex sp. 3.3
Agrostis canina 2.7
Molinia caerulea 2.3
Pteridiumapuilinum
1.5
Other 1.3
Eriophorum sp. 1.2
Agrostis capillaris 1.2
Festuca rubra 0.5
Carex sp. 0.3
Juncus sp. 0.2
Dec Other Gramineae 30
131
from till food type g/day kJ/day % of diet referencecomment
Horse Equus caballus
Dec Agrostis curtisii Putman et al, 198716
Tree leaves 13
Mosses 12
Calluna vulgaris 9.7
Ulex sp. 9
Pteridiumapuilinum
2.5
Agrostis canina 1.8
Molinia caerulea 1.5
Agrostis capillaris 1.3
Eriophorum sp. 1.2
Carex sp. 0.8
Other 0.8
Juncus sp. 0.3
Festuca rubra 0.3
Energy intake 1E+05 Putman, 1986Energy intake for New Forest ponies estimated to rangefrom 82,000 to 139,000 kJ per day.
No seasonal data
Energy intake 1E+05 Southwood et al.,1993
Racehorses kept in Australia had energy intake of121,000 kJ per day.
No seasonal data
132
from till food type g/day kJ/day % of diet referencecomment
Mink Mustela vison
Fish Chanin & Linn, 1980a% FREQUENCY in scats. a) River Teign, b) SlaptonLey freshwater ake, c) River Frome. Devon and Dorset,UK.Takes eels more frequently than does otter.34% salmonid, 17% eel
No seasonal data 54
Mammals 6% rabbit/hare, 4% bank vole, 3% field vole, 4% shrew29
Birds 4% pigeons, 4% songbirds11
Fish Chanin & Linn, 1980bSlapton Ley26% eels, 8% Cyprinid, 8% stcikleback, 6% perch,
No seasonal data 53
Mammals 11% field vole, 8% rat31
Birds 15% Ralliforms (eg moorhen), 6% ducks30
Mammals 10% rabbit/hare15
Fish Chanin & Linn, 1980cRiver Frome8% eels, 5% Salmonid, 6% Cyprinds
No seasonal data 34
Birds 16% Ralliforms (eg moorhen), 4% Ducks23
May Jul Fish Clode & Macdonald(1995)
% SPRAINTS containing food item. (Totals may exceed100%) ScotlandCommon fish included Anguillidae (28% of spraints),Gadidae (24%), Heterosomata (16%) andGasterosteidae (16%)
112
Crustacea Decapoda 24%, Isopoda 12%36
Mammals Leporidae8
Birds Haematopodidae4
133
from till food type g/day kJ/day % of diet referencecomment
Mink Mustela vison
May Jul Molluscs Clode & Macdonald(1995)
4
Insects 4
134
from till food type g/day kJ/day % of diet referencecomment
Mole Talpa europea
Coleoptera larvae Castien & Gosalbez,1995
% VOLUME in digestive tract. Spain.No seasonal data 28
Diptera larvae 22
Lumbricidae 21
Chilopoda 9
Hymenoptera 6
Adult coleoptera 3
Earthworms 30 Gorman & Stone,1990
Average free living mole will eat 60 g of food per day(freshweight) of which at least half is worms.
No seasonal data 50
Jun Aug Earthworms Stomach contents English fens. Seasonal differencemay be due to fewer worms available in summer.
50
Dec Feb Earthworms 90
Jan Feb Maggots,mealworms,neonatal mice &earthworms
22.4 170 Hawkins & Jewell,1962
Daily food intake - WET WEIGHT. Equivalent to 0.50kcal/g body wt/day.
Mar Maggots,mealworms,neonatal mice &earthworms
23 165 Daily food intake - WET WEIGHT. Equivalent to 0.42kcal/g body wt/day.
Nov Dec Maggots,mealworms,neonatal mice &earthworms
26.3 204 Daily food intake - WET WEIGHT. Laboratory study ofcaptive animals. Equivalent to 0.64 kcal/g body wt/day.
135
from till food type g/day kJ/day % of diet referencecomment
Mole Talpa europea
Earthworms 50 185 HBMDaily energy requirements for 80 g mole.No seasonal data
136
from till food type g/day kJ/day % of diet referencecomment
Muntjac Muntiacus reevesi
Bramble &raspberry
HBMResults of 2 year study in Suffolk in which Rubusspecies comprise 30 - 40% of total diet in all months.
No seasonal data 30
Bramble (Rubusspp.)
Jackson et al, 1977Data given as % VOLUME of fragments found inrumens of 26 individuals. UK. Animals killed mainly inlate winter. Frequency of occurrence = 100%.
No seasonal data 46
Other herbs Occurred in 96% of samples.11
Grasses Occurred in 85% of samples.Predominantly small sweetspecies such as Agrostis tenuis.
9.9
Bluebell Occurred in 81% of samples.8.2
Conifers, live Occurred in 27% of samples.Comprised Douglas fir(Pseudotsuga menziesii), Scots Pine (Pinus sylvestris),Norway spruce (Picea abies), Sitka spruce (Piceasitchensis).
3.4
Ivy (Hedera helix) Occurred in 12% of samples.2.2
Broad leavedtrees, live
Occurred in 27% of samples.1.2
Ranunculus spp. Occurred in 23% of samples.0.8
Rhododendronspp.
Occurred in 8% of samples.0.6
Fungi Occurred in 19% of samples.0.6
Conifers, deadneedles
Occurred in 15% of samples.0.2
Ferns, dead Occurred in 19% of samples.0.2
Betula bark Occurred in 4% of samples.0.1
Mosses Occurred in 4% of samples.0.1
May Dog's mercury Occurred in 54% of samples.15
137
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
Fish & amphibians Breathnach & Fairley,1993
Crayfish are minor food in winter, whilst consumption offish peaked in summer. Frogs eaten least in summerand more in winter and spring. Roach most importantfish in diet. Analysis of spraints, Ireland.
No seasonal data
Jan Fish Bueno, 1996% SPRAINTS containing food items.(Totals for a givenmonth may exceed 100). Spain.
93
Crayfish(Procambarusclarkii)
30
Apr Mar Salmonid Chanin, 1981a% FREQUENCY of items as a proportion of totalnumber found in all spraints. UK. (a) = River Teign, (b)= Slapton Ley.
60.1
Eel 29.5
Loach 2.3
Earthworm 2
Lagomorph 1.7
Columbiform 0.8
Rat 0.8
Amphibia 0.6
Bank vole 0.6
Field vole 0.6
Anseriform 0.3
Woodmouse 0.3
Mustelid 0.3
138
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
Apr Mar Arthropod Chanin, 1981a0.3
May Apr Cyprinid Chanin, 1981bSlapton Ley.32.1
Perch 28.2
Eel 23.2
Pike 6.3
Unidentified bird 3.3
Anseriform 3
Stickleback 1.5
Ralliform 0.7
Lagomorph 0.7
Amphibia 0.5
Unidentifiedmammal
0.2
Salmonid 0.2
Passeriform 0.2
Woodmouse 0.2
May Jul Anguillidae Clode & MacDonald,1995a
% SPRAINTS containing food item. (Totals may exceed100%) Scotland.(a) population allopatric with mink (b) populationsympatric with mink with abundant mammal and avianprey (c) population sympatric with mink with fewermammalian and avian prey species.
56
Haematopodidae 12
139
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
May Jul Gadidae Clode & MacDonald,1995a
12
Pholidae 8
Leporidae 8
Laridae/Sternidae 8
Gasterosteidae 8
Zoarcidae 4
Alcidae 4
Cottidae 4
Decapoda 4
Scolopacidae 4
Isopoda 4
Heterosomata 4
May Jul Anguillidae Clode & MacDonald,1995b
52
Vegetation 36
Gadidae 15
Decapoda 11
Ammodytidae 11
Isopoda 11
Gobiidae 7.4
Gasterosteidae 3.7
Heterosomata 3.7
140
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
May Jul Liparidae Clode & MacDonald,1995b
3.7
Stichaeidae 3.7
May Jul Isopoda Clode & MacDonald,1995c
40
Anguillidae 28
Ammodytidae 24
Vegetation 19
Gadidae 16
Decapoda 12
Gasterosteidae 8
Gobiidae 8
Cottidae 4
Stichaeidae 4
Pholidae 4
Insects Foster & Turner, 1991% SPRAINTS containing insects. Insects are activelypredated, especially water beetles of the generaHydrous and Dytiscus and the orthopteran Gryllotalpa.UK.
No seasonal data 72
Experimental diet HBMDaily food consumption of adult males is approx. 12 -15% body weight.
No seasonal data
141
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
Sep Nov Fish Jurajda et al, 1996% VOLUME of prey items in spraints. Czech Republic.Most fish taken were smaller than 200mm in length.
62
Mammals 26
Birds 12
Fish Kemenes & Nechay,1990
% FREQUENCY in spraints. Proportion of fish in dietgreater than 80%. No preference for certain species butsmall-medium size most common. Hungary.
No seasonal data 80
Aug Aug Fish 1512 6696 Nolet & Kruuk (1994)A 5.4 kg lactating female was radio-tagged andintensively observed for 19 days on Shetland.
Dec Feb Amphibians Sulkava, 1996% VOLUME in spraints. Finland. Small fish mostimportant - body length less than 15cm. Perches, pikesand cyprinids important in all seasons. Toads alsoeaten.
33
Sculpins (small) 10
Burbots 9
Mar May Cyprinids Sullkava, 199639
Jul Jun Butterfish (Pholisgunellus)
Watt, 1995a% FREQUENCY as a proportion of total number ofitems indentified in all spraints. 2 year study from twocoastal habitats in Mull. (a) = gentle slope, pebbles; (b)= steep and rocky. UK.
26
Cottids (Cottidae) 11
142
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
Jul Jun Crab (Carcinusmaenas)
Watt, 1995a9.6
Yarrell's blenny(Chirolophisascanii)
7.9
Eel (Anguillaanguilla)
7.4
Stickleback(Spinachiaspinachia)
6.4
Clingfish(Lepadogasterlepadogaster)
6
Rocklings (Ciliataspp.,Gaidropsarus spp)
3.5
Non-rocklinggadoids (Gadidae)
3.1
Gobies 3
Wrasses(Labridae)
2.4
Pipefish(Syngnathidae)
2.4
Flatfish(Pleuronectidae,Bothidae)
2.4
Shanny(Lipophrys pholis)
2.1
143
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
Jul Jun Dragonet(Callionymus lyra)
Watt, 1995a1.2
Velvet crab(Portunus puber)
1.2
Birds 1
Amphibians 0.9
Lumpsucker(Cyclopteruslumpus)
0.9
Squat lobster(Galathea spp.)
0.5
Sea snails(Liparidae)
0.5
Salmonidae 0.2
Jul Jun Butterfish (Pholisgunnellus)
Watt, 1995b25
Rocklings (Ciliataspp.,Gaidropsarusspp.)
12
Cottids (Cottidae) 12
Eel (Anguillaanguilla)
7.2
Non-rocklinggadoids (Gadidae)
6.4
144
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
Jul Jun Stickleback(Spinachiaspinachia)
Watt, 1995b5.6
Sea snails(Liparidae)
4.8
Flatfish(Pleuronectidae,Bothidae)
3.9
Crab (Carcinusmaenas)
3
Shanny(Lipophrys pholis)
2.8
Wrasses(Labridae)
2.5
Yarrell's blenny(Chirolophisascanii)
2.4
Pipefish(Syngnathidae)
2.3
Gobies 2.2
Dragonet(Callionymus lyra)
1.7
Lumpsucker(Cyclopteruslumpus)
1.3
Velvet crab(Portunus puber)
1
Gurnards(Triglidae)
0.7
145
from till food type g/day kJ/day % of diet referencecomment
Otter Lutra lutra
Jul Jun Birds Watt, 1995b0.6
Amphibians 0.5
Clingfish(Lepadogasterlepadogaster)
0.2
Conger eel(Conger conger)
0.2
146
from till food type g/day kJ/day % of diet referencecomment
Pipistrelle Pipistrellus pipistrellus
mealworms 3 HBMCaptive dietNo seasonal data
Oct Nov Diptera Hoare, 1991% FREQUENCY in faeces of bats roosting in a church,Kent.Cecidomyiidae 37.9%, Chironomidae 16.7%,Trichoceridae 15.2%, Culicidae 10.6%.
98
Lepidoptera Pyralidae2
29.3 Nagy, 1994Field Metabolic rate for a 7.3g individualNo seasonal data
Jun Aug Diptera Swift et al, 1985% FREQUENCY in faeces of 112 bats during summerin Scottish ripariam woodland.56.4% Chironomidae (non-biting midges)
69
Trichoptera(caddis flies)
23.4% Glossomatidae26
147
from till food type g/day kJ/day % of diet referencecomment
Polecat Mustela putorius
Lagomorphs HBM% VOLUME in faeces. Based on 558 scats from 20polecats in Wales.
No seasonal data 37
Wood mice 15
Bank vole 10.7
Galliformes 7.6
Other mammals 6.9
Amphibians 6
Field vole 5.7
Other birds 5.2
Passeriformes 4.2
Columbiformes 2.3
148
from till food type g/day kJ/day % of diet referencecomment
Pygmy shrew Sorex minutus
Jun Aug Araneids Churchfield, 1994% FAECES containing items. (Totals may exceed 100.)Study of shrews inhabiting watecress beds. UK.
64
Coleopteran adults 52
Isopods 48
Dipteran adults 40
Hemipteran adults 28
Dec Feb Acarines 80
Isopods 60
Araneids 50
Coleopteran adults 40
Collembolans 20
Coleopteranlarvae
20
Hemipteran adults 20
Dipteran adults 20
Lepidopteranlarvae
20
52 HBMWill eat 1.25 times bodyweight per day in captivity or 9.7-13.0kJ/g bodyweight per day.
No seasonal data
8.12 Pernetta, 1976Cites 3 studies indicating that pygny shrews willconsume 1.5, 1.85 and 2.03g of food per g bodyweight
No seasonal data
Jan Mar Coleoptera adults % Wet weight in stomach contents. Oxfordhiregrassland.
48
149
from till food type g/day kJ/day % of diet referencecomment
Pygmy shrew Sorex minutus
Jan Mar Coleoptera larvae Pernetta, 197626
Opiliones 15
Aranae 7
Apr Jun Coleoptera adults 34
Aranae 24
Coleoptera larvae 22
Opiliones 11
Jul Sep Coleoptera adults 39
Opiliones 27
Aranae 23
Coleoptera larvae 6
Oct Dec Coleoptera adults 47
Opiliones 44
Aranae 5
Coleoptera larvae 2
150
from till food type g/day kJ/day % of diet referencecomment
Rabbit Oryctolagus cuniculus
Jun Aug Dicotyledons,especially flowers
Chapuis & Gaudin,1995
Identification of plant fragments in faecal pellets.France. Aphyllanthes monspeliensis, Rubia peregrina,Thymus vulgaris and Brachypodium retusum alsoimportant.
Oct May Graminae
Carrots 453 Cowan,1987Data on carrot bait consumption by wild rabbits at 10sites in Southern England. Average consumption per kgrabbit was 286g.
No seasonal data
Dried & pelletedrye grass (cvMelle)
83 Monk, 198415 farmland rabbits, average wt 1310g fed in thelaboratory for 7-14 days. Wet weight equivalent wouldbe approx 415g.
No seasonal data
Sep Sep Mixed grassesand herbs
870 10 farmland rabbits foraged in a 24x26m grassenclosure for 22 days. The amount of grass left at theend of the experiment was compared with that in similarrabbit proof exclosures. The difference in biomassimplies that rabbits ate an average of 174g dry matter aday (136g per kg bodyweight). Assuming 80%moisture, daily wet weight consumption of grass will be870g
Jan Jan Winter barley Monk, 1984b% VOLUME in faeces collected fromHampshire.farmland.
87
Feb Feb Winter barley 84
Apr Apr Winter barley 67
May May Winter barley 75
Jun Jun Winter barley 75
151
from till food type g/day kJ/day % of diet referencecomment
Rabbit Oryctolagus cuniculus
Jul Jul Winter barley Monk, 1984bCrop harvested34
Sep Sep Winter barley New crop sown23
Dec Dec Winter barley 88
Jan Festuca Williams et al, 1974% VOLUME based on epidermal fragments of majorspecies in rabbit faeces. UK.
38
Poa trivialis 18
Others Minor species contributing to the diet were: Angelicasylvestris, Cirsium palustre, Conium maculatum,Glechoma hederacea, Hieraceum umbellatum,Hypericum perforatum, Plantago lanceolata,Ranunculus bulbosus and Urtica dioica.
15
Anthoxanthum 12
Agropyron 8
Calamagrostis 4
Dicotyledons 3
Arrhenatherum 2
Feb Festuca 50
Others 15
Agropyron 10
Poa trivialis 10
Anthoxanthum 8
Calamagrostis 6
Arrhenatherum 1
Mar Calamagrostis 23
152
from till food type g/day kJ/day % of diet referencecomment
Rabbit Oryctolagus cuniculus
Mar Festuca Williams et al, 197422
Others 18
Agropyron 13
Poa trivialis 10
Arrhenatherum 8
Anthoxanthum 5
Dicotyledons 1
Apr Festuca 38
Calamagrostis 20
Others 13
Poa trivialis 10
Arrhenatherum 8
Agropyron 6
Agropyron 5
Anthoxanthum 4
Dicotyledons 2
May Festuca 38
Others 20
Calamagrostis 16
Anthoxanthum 10
Poa trivialis 10
Agropyron 4
Arrhenatherum 2
Jun Others 35
153
from till food type g/day kJ/day % of diet referencecomment
Rabbit Oryctolagus cuniculus
Jun Calamagrostis Williams et al, 197420
Anthoxanthum 11
Festuca 10
Dicotyledons 8
Poa trivialis 7
Arrhenatherum 6
Agropyron 3
Jul Others 45
Festuca 40
Calamagrostis 6
Dicotyledons 2
Poa trivialis 2
Agropyron 2
Arrhenatherum 1
Aug Festuca 21
Calamagrostis 20
Others 17
Poa trivialis 12
Agropyron 11
Arrhenatherum 9
Anthoxanthum 7
Dicotyledons 3
Sep Festuca 48
Others 17
154
from till food type g/day kJ/day % of diet referencecomment
Rabbit Oryctolagus cuniculus
Sep Calamagrostis Williams et al, 197410
Agropyron 9
Dicotyledons 7
Anthoxanthum 5
Arrhenatherum 2
Poa trivialis 2
Oct Festuca 41
Dicotyledons 18
Dicotyledons 15
Agropyron 13
Calamagrostis 11
Others 10
Poa trivialis 5
Anthoxanthum 2
Arrhenatherum 2
Nov Festuca 43
Agropyron 18
Agropyron 17
Others 10
Poa trivialis 10
Others 9
Calamagrostis 8
Dicotyledons 5
Anthoxanthum 5
155
from till food type g/day kJ/day % of diet referencecomment
Rabbit Oryctolagus cuniculus
Nov Arrhenatherum Williams et al, 19743
Dec Festuca 45
Others 18
Poa trivialis 13
Calamagrostis 6
Agropyron 6
Dicotyledons 5
Anthoxanthum 4
Arrhenatherum 3
Coastal grasses Wolfe et al., 1996In an Irish coastal grassland system, grassesconstituted 85% of annual diet.
No seasonal data 85
156
from till food type g/day kJ/day % of diet referencecomment
Red squirrel Sciurus vulgaris
Tree seeds, fruits,berries & fungi.
HBMPrimary foods. Secondary foods include buds, shootsand flowers of trees, bark, invertebrates and lichen.Standing water may be sought in hot weather.
No seasonal data
Cedar nuts Moller, 1983% VOLUME in stomach contents. Russian study from1932 after good cedar crop.
No seasonal data 83
Fungi 16
Other conifer seed 1.3
Jan Dec Conifer seed Tittensor, 1970% STOMACHS containing food item. Conifer wood, UK.(Total values for any given season may exceed 100%).(From Moller, 1983.)
100
Conifer buds 80
Fungi 50
Conifer shoots 40
Conifer flowers 12
Mar May Conifer buds Spring.100
Conifer seed 100
Fungi 45
Conifer shoots 40
Conifer flowers 15
Jun Aug Conifer seed Summer.100
Conifer buds 55
Fungi 50
Conifer shoots 28
157
from till food type g/day kJ/day % of diet referencecomment
Red squirrel Sciurus vulgaris
Jun Aug Conifer flowers Tittensor, 197012
Sep Nov Conifer seed Autumn.100
Fungi 85
Conifer buds 50
Conifer shoots 15
Dec Feb Conifer seed Winter.100
Conifer buds 90
Conifer shoots 55
Fungi 25
Jan Dec Spruce seed Zwahlen, 1975% OBSERVATIONS of foods eaten within 1 hourperiods. Mixed conifer and deciduous. Switzerland.(From Moller, 1983).
31
Pine seed 20
Beech nut 19
Spruce buds 7
Mar May Pine seed Spring.22
Spruce seed 20
Spruce buds 12
Jun Aug Spruce seed Summer.36
Beech nut 30
Pine seed 23
Sep Nov Beech nut Autumn.29
Spruce seed 28
158
from till food type g/day kJ/day % of diet referencecomment
Red squirrel Sciurus vulgaris
Sep Nov Pine seed Zwahlen, 197516
Spruce buds 5
Dec Feb Spruce seed Winter.45
Pine seed 28
Spruce buds 16
159
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Jun Aug Cereal grain Cibien et al, 199519
Sugar beet leaves 17
Pea seeds 14
Cereal leaves 14
Pea leaves 12
Non-cultivatedplants
8
Sep Nov Cereal shoots 25
Sugar beet roots 20
Bramble 20
Tree and shrubbrowse
15
Nov Jan Cereal shoots 56
Brambles 12
Ivy 8
Sugar beet roots % DRY WEIGHT in rumen contents of 85 individuals inPicardie, northern France. Arable land, mainly cerealsand sugar beet.
5
Browse HBMIn S and E England, brambles (Rubus spp.) especiallyselected throughout year, with deciduous browse andforbs more important in summer and ivy, conifers, fernsand dwarf shrubs important in winter.
No seasonal data
Jan Rubus fruticosus Hearney & Jennings,1983
% FAECAL PELLETS containing food item. (Totals forany month may exceed 100). UK.
100
160
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Jan Festuca ovina Hearney & Jennings,1983
90
Agrostis tenuis 29
Dactylis glomerata 28
Urtica dioica 27
Holcus lanatus 11
Phleum pratense 11
Lolium perenne 11
Cynosuruscristatus
9
Poa spp. 8
Pseudotsugamenziesii
6
Hedera helix 5
Pinussylvestris/nigra
4
Quercus robur 3
Other herbs 2
Crataegusmonogyna
1
Deschampsiaflexuosa
1
Feb Rubus fruticosus 100
Festuca ovina 49
Urtica dioica 32
161
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Feb Cynosuruscristatus
Hearney & Jennings,1983
32
Agrostis tenuis 31
Lolium perenne 18
Deschampsiaflexuosa
11
Pseudotsugamenziesii
9
Pinussylvestris/nigra
7
Poa spp. 3
Phleum pratense 2
Other herbs 2
Hedera helix 2
Quercus robur 2
Filipendula ulmaria 1
Rosa spp. 1
Crataegusmonogyna
1
Mar Rubus fruticosus 98
Festuca ovina 88
Urtica dioica 55
Agrostis tenuis 43
Dactylis glomerata 21
Cynosuruscristatus
17
162
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Mar Lolium perenne Hearney & Jennings,1983
13
Deschampsiaflexuosa
9
Holcus lanatus 8
Pseudotsugamenziesii
8
Phleum pratense 6
Other herbs 4
Pinussylvestris/nigra
3
Quercus robur 3
Poa spp. 3
Rosa spp. 3
Filipendula ulmaria 1
Juncus spp. 1
Hedera helix 1
Fraxinus excelsior 1
Crataegusmonogyna
Apr Rubus fruticosus 94
Urtica dioica 80
Festuca ovina 79
Agrostis tenuis 62
Lolium perenne 17
163
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Apr Pseudotsugamenziesii
Hearney & Jennings,1983
5
Quercus robur 3
Phleum pratense 3
Other herbs 3
Crataegusmonogyna
3
Deschampsiaflexuosa
3
Holcus lanatus 3
Pinussylvestris/nigra
2
Juncus spp. 2
Rosa spp. 1
May Urtica dioica 93
Rubus fruticosus 89
Festuca ovina 83
Agrostis tenuis 51
Betula spp. 16
Holcus lanatus 13
Pseudotsugamenziesii
5
Quercus robur 5
Fraxinus excelsior 4
Deschampsiaflexuosa
3
164
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
May Filipendula ulmaria Hearney & Jennings,1983
2
Juncus spp. 2
Lolium perenne 2
Other herbs 2
Rosa spp. 1
Crataegusmonogyna
1
Poa spp. 1
Jun Urtica dioica 96
Rubus fruticosus 93
Festuca ovina 81
Agrostis tenuis 47
Betula spp. 13
Quercus robur 13
Fraxinus excelsior 5
Other herbs 4
Pseudotsugamenziesii
2
Deschampsiaflexuosa
1
Juncus spp. 1
Filipendula ulmaria 1
Lolium perenne 1
Jul Urtica dioica 100
165
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Jul Rubus fruticosus Hearney & Jennings,1983
100
Festuca ovina 76
Agrostis tenuis 52
Betula spp. 8
Quercus robur 6
Other herbs 4
Filipendula ulmaria 3
Fraxinus excelsior 3
Poa spp. 3
Pseudotsugamenziesii
2
Cynosuruscristatus
2
Pinussylvestris/nigra
1
Dactylis glomerata 1
Crataegusmonogyna
1
Aug Urtica dioica 97
Rubus fruticosus 89
Agrostis tenuis 61
Festuca ovina 59
Betula spp. 7
Filipendula ulmaria 4
166
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Aug Holcus lanatus Hearney & Jennings,1983
4
Quercus robur 3
Other herbs 3
Pseudotsugamenziesii
3
Rosa spp. 2
Poa spp. 2
Pinussylvestris/nigra
1
Crataegusmonogyna
1
Hedera helix 1
Sep Rubus fruticosus 91
Urtica dioica 90
Festuca ovina 64
Agrostis tenuis 48
Holcus lanatus 18
Poa spp. 11
Other herbs 5
Quercus robur 4
Pseudotsugamenziesii
3
Betula spp. 2
Crataegusmonogyna
1
167
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Sep Rosa spp. Hearney & Jennings,1983
1
Pinussylvestris/nigra
1
Filipendula ulmaria 1
Juncus spp. 1
Oct Rubus fruticosus 99
Urtica dioica 87
Festuca ovina 60
Holcus lanatus 23
Agrostis tenuis 21
Poa spp. 7
Pseudotsugamenziesii
7
Juncus spp. 3
Betula spp. 3
Pinussylvestris/nigra
2
Quercus robur 2
Other herbs 1
Rosa spp. 1
Deschampsiaflexuosa
1
Nov Rubus fruticosus 84
Festuca ovina 71
168
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Nov Urtica dioica Hearney & Jennings,1983
51
Holcus lanatus 21
Agrostis tenuis 18
Quercus robur 5
Pinussylvestris/nigra
5
Poa spp. 3
Pseudotsugamenziesii
3
Juncus spp. 2
Hedera helix 2
Rosa spp. 2
Lolium perenne 2
Filipendula ulmaria 1
Other herbs 1
Crataegusmonogyna
1
Cynosuruscristatus
1
Dec Rubus fruticosus 99
Festuca ovina 83
Urtica dioica 36
Agrostis tenuis 25
Holcus lanatus 14
169
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Dec Hedera helix Hearney & Jennings,1983
8
Pinussylvestris/nigra
5
Pseudotsugamenziesii
4
Quercus robur 3
Dactylis glomerata 3
Other herbs 3
Juncus spp. 2
Crataegusmonogyna
1
Filipendula ulmaria 1
Cynosuruscristatus
1
Mar May Grasses &grasslike plants
Henry, 1978% VOLUME in rumen. UK.28
Heather 25
Bilberry 12
Other forbs 12
Rosebay willow-herb
7
Sitka spruce 6
Scots pine 6
170
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Mar May Other dwarfshrubs
Henry, 19782
Ferns 1
Other trees 1
Jun Aug Heather Summer.27
Rosebay willow-herb
24
Sitka spruce 18
Other forbs 9
Grasses &grasslike plants
8
Grasses andgrasslike plants
8
Douglas fir 7
Fungi 4
Bilberry 2
Sep Nov Heather Autumn.42
Bramble & gorse 14
Other trees 11
Grasses &grasslike plants
9
Scots pine 8
Ferns 6
Bilberry 3
Fungi 3
171
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Sep Nov Other forbs Henry, 19781
Sitka spruce 1
Dec Feb Heather Winter.41
Scots pine 28
Grasses &grasslike plants
11
Sitka spruce 9
Bilberry 3
Other dwarfshrubs
3
Other forbs 2
Bramble & gorse 2
Jan Conifers Jackson, 1980% VOLUME in rumen analysis. New Forest, UK.33
Bramble/rose 31
Ivy 12
Other 6
Heather 6
Herbs 5
Grasses Included Agrostis tenuis, A. canina, Deschampsiaflexuosa, Poa annua, Holcus lanatus and Sieglingiadecumbens. Appeared to avoid Agrostis setacea,Molinia caerulea, Deschampsia cespitosa andBrachypodium sylvaticum.
4
Other broadleaves 2
172
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Jan Fruits Jackson, 19801
Feb Bramble/rose 26
Conifers 22
Ivy 22
Heather 14
Grasses 5
Other broadleaves 4
Herbs 2
Other 2
Holly 2
Fruits 1
Mar Bramble/rose 26
Conifers 22
Ivy 22
Heather 14
Grasses 5
Other broadleaves 4
Other 3
Herbs 2
Fruits 1
Apr Herbs 30
Bramble/rose 20
Heather 14
Other broadleaves 14
173
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Apr Grasses Jackson, 198010
Ivy 7
Conifers 5
May Bramble/rose 35
Other broadleaves 30
Herbs 16
Grasses 7
Ivy 6
Heather 5
Conifers 1
Jun Bramble/rose 32
Herbs 16
Other broadleaves 14
Holly 14
Conifers 8
Grasses 8
Heather 4
Ivy 4
Jul Bramble/rose 40
Herbs 16
Other broadleaves 15
Other 10
Grasses 8
Heather 7
174
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Jul Ivy Jackson, 19803
Holly 1
Aug Bramble/rose 40
Herbs 17
Other broadleaves 15
Other 10
Grasses 8
Heather 7
Ivy 3
Sep Bramble/rose 38
Herbs 17
Other broadleaves 14
Fruits 8
Grasses 8
Other 7
Heather 5
Ivy 3
Oct Bramble/rose 38
Conifers 12
Ivy 11
Grasses 9
Other 8
Fruits 8
Heather 5
175
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Oct Other broadleaves Jackson, 19805
Herbs 4
Nov Bramble/rose 37
Fruits 17
Conifers 12
Ivy 11
Grasses 10
Other broadleaves 5
Heather 4
Herbs 4
Dec Bramble/rose 46
Other 13
Conifers 13
Heather 7
Fruits 7
Herbs 6
Grasses 4
Ivy 2
Other broadleaves 2
Heather Jong et al., 1995A positive selection was found for heather, otherdicotyledons and evergreen ferns which togetherformed bulk of diet. Coarse monocotyledons wereavoided. Sitka spruce formed a significant part of thediet in spring. Study in N. England.
No seasonal data
176
from till food type g/day kJ/day % of diet referencecomment
Roe deer Capreolus capreolus
Arable Putman, 1986% FREQUENCY animals seen in habitat on standardtransects. Two sites in S. Hampshire. UKGrazing on arable land most commonly seen betweenApril and June. Highest figure was 55% for wheat inApril.
No seasonal data 55
oilseed rape Reynolds & Tapper,1990
Wild Roe deer in Scotland. Deer found to spendconsiderable time feeding in oilseed rape fields (36% ofthe time away from cover). Although it is an importantfood to the deer, the winter of study was mild and moretime would be expected on brassicae crops duringcolder weather.
No seasonal data
Captive diet Szmidt, 1975Captive animals offered a variety of browse from foresttrees and shrubs. Poland.In winter, preferred species were Fagus silvatica,Evonymus europea and Carpinus betulus. In summer,preferred species were Quercus robur, Evonymuseuropea, Sorbus aucuparia and Salix sp.
No seasonal data
177
from till food type g/day kJ/day % of diet referencecomment
Stoat Mustela erminea
Birds/eggs Day, 1963a% FREQUENCY in gut. UK. (From Handbook of BritishMammals).
No seasonal data 37
Lagomorphs 32
Small rodents 26
Water voles & rats 3
Insectivores 2
Birds/eggs Day, 1963bIreland.No seasonal data 38
Lagomorphs 35
Small rodents 18
Water voles & rats 3
Birds Day, 1968% FREQUENCY from gut analysis. UK.Also take earthworms, rats, large insects and theirlarvae.
No seasonal data 33
Lagomorphs 28
Small rodents 22
Lagomorphs Fairley, 1971% FREQUENCY in gut contents. Ireland.No seasonal data 40
Birds 38
Small rodents 17
Food requirement 57 HBM57 g freshweight required by males, 33 g freshweightrequired by females.
No seasonal data
178
from till food type g/day kJ/day % of diet referencecomment
Stoat Mustela erminea
Rabbit HBMBritish Isles study found rabbit constitutes one third ofdiet, supplemented with small rodents, brown rats,squirrels, birds and others.
No seasonal data 33
King, 1989Road killed Stoats from S. Ireland. 29 rabbits, 24shrews, 17 birds, 13 rats, 5 wood mice and 3 bankvoles, (rare in Ireland cf. England)
No seasonal data
Small rodents Pounds, 1981% FREQUENCY in gut and scats. Scotland, UK. (FromHandbook of British Mammals).
No seasonal data 42
Lagomorphs 38
Birds/eggs 18
179
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Natural diet Castien, 1995Identified 21 major taxonomic groups in the diet. Notaxonomic group dominated indicating generalistfeeding. Takes more aquatic prey in winter and spring.
No seasonal data
Mar May Asellus Churchfield, 1984% FAECES containing prey items. (Totals for a givenseason may exceed 100%) UK.In British water cress beds, 33 - 67% (mean 50%) preytaken is aquatic, the dominant prey in all months beingfreshwater crustaceans, (Gammarus and Asellus)
80
Trichopteranlarvae (cased)
74
Dipteran adults 49
Gastropods 29
Lumbricids 21
Plecopteran larvae 19
Geophilomorphs 18
Isopods 16
Staphylinids 14
Gammarus 13
Ostracods 13
Araneids 13
Coleoptera larvae 13
Carabids 11
Acarines 11
Other dipteranlarvae
11
180
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Mar May IndeterminateColeoptera adults
Churchfield, 198410.5
Trichopteranadults
5
Formicids 5
Lepidopteranlarvae
5
Simulium sp.larvae
5
Ephemeropteranlarvae
4.2
Coleopteran adults 4
Hemipteran adults 2.6
Lithobiomorphs 1.3
Diplopods 1.3
Trichopteranlarvae (uncased)
1.3
Other dipteranlarvae
1.3
Jun Aug Asellus 70
Dipteran adults 59
Gammarus 33
Other dipteranlarvae
29
Trichopteranlarvae (cased)
23
181
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Jun Aug Other dipteranlarvae
Churchfield, 198419
IndeterminateColeoptera adults
19
Formicids 18
Plecopteran larvae 17
Hemipteran adults 17
Isopods 16
Acarines 16
Araneids 15
Opilionids 14
Gastropods 14
Staphylinids 14
Diplopods 13
Ostracods 11
Lumbricids 11
Carabids 10.2
Simulium sp.larvae
10
Chrysomelids 8.5
Geophilomorphs 5
Hemipteran adults 4.2
Trichopteranlarvae (uncased)
3.6
Tipulid larvae 2.9
182
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Jun Aug Coleopteran adults Churchfield, 19842.9
Dermapterans 1.8
Lepidopteranlarvae
1.8
Sep Nov Asellus 63
Dipteran adults 56
Plecopteran larvae 42
Trichopteranlarvae (cased)
40
Gammarus 31
Other dipteranlarvae
29
Lumbricids 27
Gastropods 25
IndeterminateColeoptera adults
21
Simulium sp.larvae
21
Staphylinids 15
Acarines 13
Ostracods 13
Isopods 8.3
Coleoptera larvae 8.3
Formicids 6.3
Geophilomorphs 6.3
183
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Sep Nov Trichopteranlarvae (uncased)
Churchfield, 19846.3
Araneids 6.3
Trichopteranadults
4.2
Diplopods 4.2
Osteichthyes 4.2
Collembolans 4.2
Coleopteran adults 2.2
Carabids 2.1
Lepidopteranlarvae
2.1
Gastropods 2.1
Lithobiomorphs 2.1
Opilionids 2.1
Dec Feb Asellus 78
Dipteran adults 54
Lumbricids 28
Gastropods 26
Gammarus 20
Other dipteranlarvae
20
IndeterminateColeoptera adults
18
Trichopteranlarvae (cased)
18
184
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Dec Feb Araneids Churchfield, 198415
Other dipteranlarvae
13
Acarines 8.6
Hemipteran adults 7.7
Geophilomorphs 7.7
Tipulid larvae 7.7
Diplopods 6.3
Ostracods 5.4
Isopods 4.4
Ephemeropteranlarvae
4.3
Trichopteranlarvae (uncased)
4.2
Hymenopterans 3.3
Lepidopteranlarvae
3.3
Plecopteran larvae 3.3
Carabids 3.3
Coleopteran adults 2.2
Coleopteranlarvae
2.2
Staphylinids 2.2
Opilionids 2.2
Coleoptera larvae 2.1
185
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Dec Feb Lithobiomorphs Churchfield, 19842.1
Aquatic prey Churchfield, 1994% FREQUENCY in faecal samples. UK.No seasonal data 56
Epigeal prey Surface-dwelling27
Hypogeal prey Soil-dwelling18
Natural diet Dupasquier &Cantoni, 1992
Minimum of 80% of prey is taken from river. Dietsignificantly correlated with density of streaminvertebrates.
No seasonal data
Nov Jan 8.2 93 Hawkins & Jewell,1962
Wet weight. Laboratory study with captive shrews. Dailyfood intake was between 8 - 12.7g wet weight and1.11 - 1.38 kcal/g body weight.
Jun Aug Simuliidae larvae Niethammer, 1978% VOLUME of stomach contents. 45 indiviudals,Austria.
18
Ephemeropteralarvae
17
Trichoptera larvae 17
Plecoptera larvae 15
Indeterminateimagines
13
Diptera imagines 3.9
Limoniidae larvae 3.9
Chironomidaelarvae
2.4
186
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Jun Aug Opiliones Niethammer, 19781.6
Lumbricidae 1.6
Psychodidaelarvae
1.6
Soricidae 0.8
Neuropteraimagines
0.8
Coleopteraimagines
0.8
Ephemeropteraimagines
0.8
Amphipoda 0.8
Dixidae 0.8
Odonata 0.8
Sep Nov Radix peregra Wolk, 1976% FREQUENCY in food remains of prey animals left inrafts.
58
Rana temporaria 15
Nepa sp. 8.6
Planorbariuscorneus
4
Molanna sp. 3.3
Galba corvus 1.3
Viviparuscontectus
1.2
Dytiscus sp. 0.7
187
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Sep Nov Planorbisplanorbis
Wolk, 19760.7
Notidobia sp. 0.7
Segmentina nitida 0.7
Dec Feb Linnephilus sp. 38
Rana temporaria 20.2
Notidobia sp. 6.5
Nepa sp. 6
Galba corvus 4.1
IndeterminateTrichoptera
2.5
Gasterosteusaculeatus
2
Bithyniatentaculata
1.5
Planorbariuscorneus
1.3
Radix peregra 1.1
Molanna sp. 1.1
Dytiscus sp. 0.9
Viviparuscontectus
0.6
Anodonta sp. 0.6
Anabolia sp. 0.4
Lymnaea stagnalis 0.3
Pisidium sp. 0.3
188
from till food type g/day kJ/day % of diet referencecomment
Water shrew Neomys fodiens
Dec Feb Galba truncatula Wolk, 19760.1
Planorbisplanorbis
0.1
Segmentina nitida 0.1
Anisus sp. 0.1
Bufo bufo 0.1
Sorex araneus 0.1
Bathyomphaluscontortus
0.1
189
from till food type g/day kJ/day % of diet referencecomment
Water vole Arvicola terrestris
249 HBMApproximately 80% of body weight consumed daily.No seasonal data
Juveniles eat fewer reeds and rhizomes than adults.Voles may eat the whole stem of young grasses; morefrequently only the growth meristems.
Mar May Herbs Strachan & Jeffries,1993
% FREQUENCY in feeding remains. UK. Fruitsincluded in diet in autumn, especially apples. In winter,the roots and bark of trees and shrubs are eaten,together with bulbs, rhizomes and roots of herbaceousspecies. At other times of the year aerial parts of plantsare the major components and in spring, leaf and flowerbuds may be taken.
40
Grasses 21
Sedges 15
Woody plants 14
Aquatics 9.5
Invertebrates 1.6
Jun Aug Herbs 49
Grasses 19
Sedges 15
Aquatics 11
Woody plants 6.6
Invertebrates 0.6
Sep Nov Herbs 43
Woody plants 19
Grasses 18
190
from till food type g/day kJ/day % of diet referencecomment
Water vole Arvicola terrestris
Sep Nov Sedges Strachan & Jeffries,1993
13
Aquatics 6
Invertebrates 1.1
Dec Feb Woody plants 39
Herbs 20
Sedges 16
Grasses 15
Aquatics 6
Invertebrates 4.3
191
from till food type g/day kJ/day % of diet referencecomment
Weasel Mustela nivalis
Small rodents Day, 1968% FREQUENCY in gut. UK. (From Handbook of BritishMammals).
No seasonal data 55
Lagomorphs 18
Birds/eggs 16
Water voles & rats 3
Insectivores 2
Mar Aug Small rodents Erlinge, 1975% FREQUENCY in 148 scats. Sweden.Microtis agrestis 41%, Clethrionomys glareolus 2%,Apodemus sp 6%, Arvicola terrestris 10%
59
Lagomorphs 31
Birds 4
Shrews 2
Sep Feb Small rodents Microtis agrestis 48%, Clethrionomys glareolus 14%,Apodemus sp 14%, Arvicola terrestris 19%,
94
Lagomorphs 5
Jan Dec Clethrionomys King, 1980a% SCATS containing prey item.(Totals may exceed100%) UK.
41
Microtus 19
Apodemus 16
Passerines 12
Eggs 7
Unidentified smallrodents
7
192
from till food type g/day kJ/day % of diet referencecomment
Weasel Mustela nivalis
Jan Dec Unidentified voles King, 1980a7
Unidentified birds 3
Non-passerines 2
Mole 0.5
Lagomorph 0.5
Jan Feb Clethrionomys King, 1980b% SCATS containing prey item. (Totals may exceed100%) UK. Seasonal data.
42
Birds 22
Microtus 22
Apodemus 12
Mar Apr Clethrionomys 38
Microtus 18
Birds 17
Apodemus 17
Eggs 9
Mole 2
Lagomorph 2
May Jun Clethrionomys 40
Eggs 38
Birds 23
Microtus 8
Apodemus 8
Jul Aug Clethrionomys 25
193
from till food type g/day kJ/day % of diet referencecomment
Weasel Mustela nivalis
Jul Aug Microtus King, 1980b23
Apodemus 22
Eggs 8
Birds 6
Sep Oct Clethrionomys 42
Apodemus 20
Microtus 17
Birds 5
Nov Dec Clethrionomys 41
Birds 18
Apodemus 17
Microtus 17
Small rodents Moors, 1975Data is % DIET COMPOSITION. UK. Woodmicecomprised 3% of diet in spring and 33% after harvest.
No seasonal data 67
Small rodents % FREQUENCY in scats. UK. Woodmice comprised3% of diet in spring and 33% after harvest.
65
Birds with/withouteggs
18
Lagomorphs 15
Insectivores 2
Captive diet(males)
39.6 Moors, 1977Average captive diet of mice, voles, rabbit and starling.0.33g/g body weight.
No seasonal data
194
from till food type g/day kJ/day % of diet referencecomment
Weasel Mustela nivalis
Captive diet(females)
22.7 Moors, 1977Average captive diet of mice, voles, rabbit and starling.0.36g/g body weight.
No seasonal data
Microtus agrestis 32.56 269.1 Moors, 1977 fCaptive female weighing 74gNo seasonal data
Rabbit 32.3 199 Captive female weighing 85g
Rabbit 32.2 199 Captive female weighing 70g
Apodemussylavaticus
28.35 199.8 Captive female weighing 81g
Apodemussylavaticus
26.6 186.6 Captive female weighing 95g
Starling 21.84 185.8 Captive female weighing 78g
Starling 20.64 176.1 Captive fwmale weighing 86g
Microtus agrestis 51.6 425.9 Moors, 1977 mCaptive male weighing 129gNo seasonal data
Rabbit 46.62 289.7 Captive male weighing 126g
Rabbit 43.71 271.1 Captive male weighing 141g
Apodemussylavaticus
41.4 288.4 Captive male weighing 115g
Apodemussylavaticus
40.04 280.9 Captive male weighing 143g
Starling 35.96 305.8 Captive male weighing 124g
Starling 30 257.1 Captive male weighing 150g
Small rodents Pounds, 1981% FREQUENCY in gut contents and scats. Scotland.UK.
No seasonal data 65
195
from till food type g/day kJ/day % of diet referencecomment
Weasel Mustela nivalis
Lagomorphs Pounds, 1981No seasonal data 22
Birds with/withouteggs
8
Small rodents Tapper, 1979% FREQUENCY in gut contents. 687 samples from 25sq km farmland site in Sussex, UKMicrotus 37%, Clethrionomys 9%, Apodemus 5%
No seasonal data 58
Birds Songbirds 15%, gamebirds 2%, eggs 1.9%.Concentrated upon by both sexes, especially in yearswhen field voles scarce.
19
Lagomorphs Mainly MALE weasels17
Insectivores 2
Water voles & rats 2
196
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Monocotyledon(Barley) testa &endosperm
Flowerdew &Gardner, 1978
% VOLUME in stomach contents. UK.No seasonal data 53
Insect 20
Fraxinus testa &endosperm
14
Unknown 13
Monocotyledonleaf
1
Jan Plant material Gorman & Zubaid,1993a
% VOLUME in stomach contents. UK. Comparison ofdifferent habitats; (a) = woodland site; (b) = sand-dunesite.
80
Insects 10
Oligochaeta 2
Feb Plant material 70
Insects 13
Oligochaeta 8
Mar Plant material 85
Insects 15
Apr Plant material 70
Insects 17
Oligochaeta 10
May Plant material 75
Insects 15
197
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
May Oligochaeta Gorman & Zubaid,1993a
5
Jun Plant material 80
Insects 14
Jul Plant material 85
Insects 15
Aug Plant material 80
Insects 20
Sep Plant material 75
Insects 17
Oct Plant material 98
Nov Plant material 95
Dec Plant material 90
Insects 5
Jan Insects Gorman & Zubaid,1993b
68
Plant material 20
Feb Insects 45
Plant material 22
Oligochaeta 22
Mar Insects 46
Oligochaeta 25
Plant material 20
198
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Apr Insects Gorman & Zubaid,1993b
60
Oligochaeta 15
Plant material 15
May Insects 56
Oligochaeta 15
Plant material 10
Jun Insects 40
Oligochaeta 38
Plant material 6
Jul Insects 55
Oligochaeta 20
Plant material 15
Aug Insects 45
Plant material 20
Oligochaeta 15
Sep Insects 46
Plant material 22
Oligochaeta 15
Oct Insects 47
Plant material 20
Oligochaeta 15
Nov Insects 44
Plant material 30
199
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Nov Oligochaeta Gorman & Zubaid,1993b
20
Dec Insects 50
Plant material 28
Oligochaeta 12
Rose hips HBMAchenes (pips) of rose hips opened to extract seed butflesh discarded.
No seasonal data
Jan Jan Cereal grain Pelz, 1989% Volume of stomach contents from individuals caughton arable farms in the Rhineland, Germany. Data werecollected from 346 individuals trapped over a 7 yearperiod.
53
Insect Larvae Data from 21 individuals17
Dicot seeds 15
Vegetative planttissue
15
Feb Feb Vegetative planttissue
Data from 14 individuals34
Cereal grain 28
Dicot seeds 20
Earthworms 12
Insect Larvae 6
Mar Mar Insect Larvae Data from 56 individuals25
Earthworms 23
Cereal grain 23
200
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Mar Mar Vegetative planttissue
Pelz, 198922
Sugar Beet seeds 7
Apr Apr Insect larvae Data from 49 individuals45
Earthworms 26
Vegetative planttissue
24
Cereal grain 5
May May Earthworms Data from 16 individuals40
Cereal grain 30
Vegetative planttissue
16
Insect larvae 10
Dicot seeds 4
Jun Jun Cereal grain Data from 15 individuals32
Dicot seeds 25
Insect larvae 25
Earthworms 9
Vegetative planttissue
9
Jul Jul Cereal grain Data from 10 individuals48
Insect larvae 28
Dicot seeds 16
Vegetative planttissue
8
201
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Aug Aug Cereal grain Pelz, 1989Data from 41 individuals37
Insect larvae 28
Dicot seeds 20
Vegetative planttissue
10
Earthworms 5
Sep Sep Cereal grain Data from 18 individuals33
Insect larvae 25
Dicot seeds 20
Earthworms 13
Vegetative planttissue
9
Oct Oct Insect larvae 30
Cereal grain Data from 48 individuals30
Vegetative planttissue
25
Dicot seeds 15
Nov Nov Vegetative planttissue
40
Cereal grain Data from 36 individuals40
Insect larvae 9
Dicot seeds 8
Earthworms 3
Dec Dec Vegetative planttissue
40
202
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Dec Dec Cereal grain Pelz, 1989Data from 22 individuals40
Insect larvae 9
Dicot seeds 8
Earthworms 3
Sugar beet seeds Pelz, 1989aCaptive woodmice ate an average of 800 seeds (3-4mgeach) per night. The maximum consumption by oneindividual was 1400 seeds in a night
No seasonal data
Mar May Monocotyledons Rogers & Gorman,1995
% VOLUME in gut contents. Set-aside land, UK. Dietcontained 87% plant material, 65% of which was greenleaves. Much less seed and animal material were eatenon setaside than in other habitats.
72
Insects 13
Other animalmaterial
10
Dicotyledons 5
Jun Aug Monocotyledons 45
Seeds Majority of seed material was Cerastium holosteoides.42
Other plantmaterial
5
Dicotyledons 3
Other animalmaterial
2
Insects 1
Sep Nov Monocotyledons 50
203
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Sep Nov Seeds Rogers & Gorman,1995
35
Other plantmaterial
6
Insects 5
Dicotyledons 2
Other animalmaterial
1
Dec Feb Monocotyledons 68
Seeds 25
Insects 5
Other animalmaterial
1
Other plantmaterial
1
Insects & animalmaterial
Rogers, 1990% VOLUME in stomach contents. Oilseed rape fieldsUK.
No seasonal data 70
Jan Seed endosperm Watts, 1968% VOLUME in stomach contents analysis. Wythamwoods, Oxford, UK.
98
Feb Seed endosperm 99
Mar Seed endosperm 62
Dicotyledon stem 27
Dead leaf 4
Blackberry bud 3
204
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Apr Seed endosperm Watts, 196844
Blackberry bud 38
Dicotyledon stem 16
May Animal Nearly all examples were of leaf-eatrng caterpillarsleaving trees to pupate in the soil.
88
Seed endosperm 7
Dicotyledon stem 4
Jun Seed endosperm 50
Animal Mainly centipedes (Lithobius)43
Ash keys 1
Fungi 1
Dead leaf 1
Dog's mercuryseed testa
1
Dicotyledon stem 1
Jul Seed endosperm 73
Animal 15
Blackthornleaf/stem
8
Fungi 2
Other dicotyledonleaf
1
Sep Seed endosperm 30
Blackberry (fruit) 22
Elderberry (fruit) 22
205
from till food type g/day kJ/day % of diet referencecomment
Wood mouse Apodemus sylvaticus
Sep Sycamore seed Watts, 196817
Flower 8
Other dicotyledonleaf
2
Oct Seed endosperm 79
Elderberry (fruit) 15
Blackberry (fruit) 2
Bracken 1
Ash keys 1
Nov Seed endosperm 69
Sycamore seed 14
Fungi 7
Fibre fromsycamore keys
4
Hawthorn (fruit) 3
Blackberry (fruit) 1
Invertebrates Zubaid & Gorman,1991
Wood mice inhabiting dune system in Scotland fed oninvertebrates, with coleopterans forming bulk of diet.
No seasonal data 70
206
207
Bibliography
Aitchison, C. W. (1987). Winter energy requirements of soricine shrews.Mammal Rev.17:25-38
Arnold, H. R. (1993)Atlas of mammals in Britain( HMSO, London).
Aulak, W. & Babinska-Werka, J. (1990). Use of agricultural habitats by Roe deerinhabiting a small forest area.Acta theriol35: 121-128
Barnes, R. F. W., Tapper, S. C. & Williams, J.(1983). Use of pastures by brown hares.J.Appl. Ecol.20: 179-185
Biancardi, C. M., Pavesi, M. & Rinetti, L. (1993). Food of European badger (Meles melesL.) in Northern Luino area (Varese County, Italy).Atti della Societa Italiana di ScienzeNaturali e del Museo Civico di Storia Naturale di Milano134: 265-280
Boag, B., MacFarlane Smith, W. H. & Griffiths, D. W. (1990). Observations on thegrazing of double oilseed rape and other crops by Roe deer.Applied AnimalBehavioural Science28: 213-220
Bolls, N. J. & Perfect, J. R.(1972). Summer resting metabolic rate of the grey squirrel.Physiol. Zool.45: 54-59
Boone, G. C. & Tinklin, R. (1988). Importance of hedgerow structure in determining theoccurrence and density of small mammals.Aspects of Applied Biology16: 73-78
Borkowski, J. (1994). Food composition of red fox in the Tatra National Park.Acta theriol.39: 209-214
Breathnach, S. & Fairley, J. S.(1993). The diet of ottersLutra lutra (L.) in the Clare Riversystem.Biology and Environment93B: 151-158
Bueno, F.(1996). Competition between american minkMustela visonand otterLutra lutraduring winter.Acta theriol.41: 149-154
Bullock, D. J. (1985). Annual diets of hill sheep and feral goats in southern Scotland.J. Appl.Ecol.22: 423-433
Caldwell, J. F., Chapman, D. I. & Chapman, N.(1983). Observations on the autumn andwinter diet of Fallow deer (Dama dama). J. Zool.201: 559-563
Castien, E.(1995). The diet ofNeomys fodiensin the Spanish Western Pyrenees.Folia Zool.44: 297-303
208
Castien, E. & Gosalbez, J.(1995). Trophic niche characterization ofTalpa europaeaLinnaeus, 1758, in the West Pyrenees (north of Iberian Peninsula).Mammalia59: 43-50
Castien, E. & Gosalbez, J.(1996). Diet ofClethrionomys glareolusin the Western Pyrenees(North Iberian Peninsula).Folia Zool.45: 137-144
Chanin, P. (1981). The diet of the otter and its relations with the feral mink in two areas ofSouthwest England.Acta theriol.26: 83-95
Chanin, P. R. F. & Linn, I. J. (1980) The diet of feral mink (Mustela vison Schreber) inSouthwest Britain.J. Zool. , Lond.192: 205-223.
Chapman, N., Harris, S. & Stanford, A. (1994) Reeves' MuntjacMuntiacus reevesiinBritain: their history, spread, habitat selection, and the role of human intervention inaccelerating their dispersal.Mammal Rev.24: (3):113-160.
Chapuis, J. L. & Gaudin, J. C. (1995). Use of food resources by the wild rabbit(Oryctolagus cuniculus) in a managed arrid garrigue scrubland.Gibier Faune Sauvage12: 213-230
Christian, S. F. (1993). The behavioural ecology of the Eurasian badger, Meles meles: spaceuse, territoriality and social behaviour.PhD Thesis, University of Sussex
Churchfield, S. (1982). Food availability and the diet of the common shrew,Sorex araneus,in Britain. J. Anim. Ecol.51: 15-28
Churchfield, S. (1982) The influence of temperature on the activity and food consumption ofthe common shrew.Acta theriol.27: 295-304.
Churchfield, S. (1984). Dietary separation in three species of shrew inhabiting water-cressbeds.J. Zool. , Lond.204: 211-228
Churchfield, S. (1985) The feeding ecology of the European Water shrew.Mammal Rev.15:13-31.
Churchfield, S. (1990)The Natural History of Shrews( Christopher Helm Ltd, Bromley).
Churchfield, S. (1994). Foraging strategies of shrews, and the evidence from field studies.Carnegie Museum of Natural History Special Publication18: 77-87
Cibien, C., Bideau, E., Boisaubert, B., Biran, H. & Angibault, J. M. (1995). Seasonal dietand habitat use in field Roe deer (Capreolus capreolus) in the Picardie region.GibierFaune Sauvage12: 37-49
209
Clode, D. & MacDonald, D. W. (1995). Evidence for food competition between mink(Mustela vison) and otter (Lutra lutra) on Scottish islands.J. Zool. , Lond.237: 435-444
Cooke, A. S., Green, P. & Chapman, N. G.(1996). Mortality in a feral population ofmuntjac muntiacus-reevesi in england.Acta theriol.41: 277-286
Corbet, G. B. & Harris, S. (1991)The Handbook of British Mammals( BlackwellScientific, Oxford).
Cowan, D. P., Vaughan, J. A. & Christer, W. G.(1987) Bait consumption by the Europeanrabbit in southern England.J. Wildl. Manage.51: (2):386-392.
Cowan, D. P., Hardy, A. R., Vaughan, J. P. & Christie, W. G.(1989) Rabbit rangingbehaviour and its implications for the management of rabbit populations. inMammalsas Pests(ed Putman, R.J.) 178-185 ( Chapman and Hall, London).
Day, M. G. (1963). An ecological study of the stoat (Mustela erminea) and the weasel(mustela nivalis) with particular reference to their food and feeding habits. University ofExeter.
Day, M. G. (1968) Food habits of British stoats (Mustela ermineaL) and weasels (Mustelanivalis L). J. Zool. , Lond.155: 485-487.
Dell'Omo, G. & Shore, R. F. (1996). Behavioral effects of acute sublethal exposure todimethoate on wood mice, Apodemus sylvaticus: II- Field studies on radio-tagged micein a cereal ecosystem.Arch. Environ. Contam. Toxicol.31: 538-542
Dickman, C. R. (1986). Habitat utilization and diet of the harvest mouse,Micromys minutus,in an urban environment.Acta theriol.31: 249-256
Dupasquier, A. & Cantoni, D. (1992). Shifts in benthic macroinvertebrate community andfood habits of the water shrewNeomys fodiensSoricidae: Insectivora.Acta oecol.13:81-99
Edgar, W. H. & Isaacson, A. J.(1974) Observations on skylark damage to sugar beet andlettuce seedlings in East Anglia.Annals of Applied Biology76: 335-337.
Eldridge, J. (1971). Some observations on the dispersion of small mammals in hedgerows.J.Zool.165: 530-534
Erlinge, S. (1975). Feeding habits of the weaselMustela nivalisin relation to preyabundance.Oikos26: 378-384
Evans, D. M. (1973). Seasonal variations in the body composition and nutrition of the vole,Microtus agrestis. J. Anim. Ecol.42: 1-18
210
Faber, J. & Ma, W. (1986). Observations on seasonal dynamics in diet composition of thefield vole,Microtus agrestis, with some methodological remarks.Acta theriol.31: 479-490
Fairley, J. S. (1971). New data on the Irish stoat.Irish Nat. J.17: 49-57
Ferns, P. N.(1976). Diet of a Microtus agrestis population in south west Britain.Oikos27:506-511
Ferrari, N. & Weber, J. M. (1995). Influence of the abundance of food resources on thefeeding habits of the red fox,Vulpes vulpes, in western Switzerland.J. Zool. , Lond.236: 117-129
Fitzgibbon, C. D. (1997). Small mammals in farm woodlands: The effects of habitat,isolation and surrounding land-use patterns.J. Appl. Ecol.34: 530-539
Flowerdew, J. R.(1997). Mammal biodiversity in agricultural habitats.1997 BCPCSymposium Proceedings no. 6969: 25-40
Flowerdew, J. R. & Gardner, G. (1978). Small rodent populations and food supply in aDerbyshire ashwood.J. Anim. Ecol.47: 725-740
Foster, J. & Turner, C. (1991). Insects and otter diet.Entomologist110: 166-169
Frylestam, B. (1986). Agricultural land use effects on the winter diet of Brown Hares (LepuseuropaeusPallas) in southern Sweden.Mammal Rev.16: 157-161
Gorecki, A. (1971). Metabolism and energy budget in the harvest mouse.Acta theriol.16:213-220
Gorman, M. L. & Stone, R. D. (1990)The Natural History of Moles( Christopher HelmLtd, Bromley).
Gorman, M. L. & Zubaid, A. (1993). A comparative study of the ecology of woodmiceApodemus sylvaticusin two contrasting habitats: deciduous woodland and maritimesand dunes.J. Zool. , Lond.229: 385-396
Goszczynski, J.(1986). Diet of foxes and martens in Central Poland.Acta theriol.31: 491-506
Green, R.(1979). The ecology of wood mice (Apodemus sylvaticus) on arable farmland.J.Zool.118: 357-377
211
Guillet, C., Bergstrom, R., Cederlund, G., Bergstrom, J. & Ballon, P.(1995). Comparisonof telemetry and pellet-group counts for determining habitat selectivity by Roe deer(Capreolus capreolus) in winter.Gibier Faune Sauvage12: 253-269
Hansson, L. (1971) Small rodent food, feeding and population dynamics. A comparisonbetween granivorous and herbivorous species in Scandinavia.Oikos22:183-198,.
Hansson, L.(1985). The food of Bank voles, Wood mice and Yellow-Necked mice.Symp.zool. Soc. Lond.55: 141-168
Harris, S. (1981). The food of suburban foxes (Vulpes vulpes), with special reference toLondon.Mammal Rev.11: 151-168
Harris, S. & Woollard, T. (1990) The dispersal of mammals in agricultural habitats inBritain. in Species Dispersal in Agricultural Habitats(eds Bunce, R.G.H. & Howard,D.C.) 159-188 ( Belhaven Press, Institute of Terrestrial Ecology, London).
Harris, S. Morris, P. Wray, S. & Yalden, D. (1995)A review of British mammals:population estimates and conservation status of British mammals other than Cetaceans( JNCC, Peterborough).
Hawkins, A. E. & Jewell, P. A. (1962). Food consumption and energy requirements ofcaptive British shrews and the mole.Proc. zool. Soc. Lond.138: 137-155
Hearney, A. W. & Jennings, T. J.(1983). Annual foods of the Red deer (Cervus elaphus)and the Roe deer (Capreolus capreolus) in the east of England.J. Zool.201: 565-570
Henry, B. A. M. (1978). Diet of Roe deer in an English conifer forest.J. Wildl. Manage.42:937-940
Hoare, L. R. (1991) The diet ofPipistrellus pipistrellusduring the pre-hibernal period.J.Zool. , Lond.225: 665-670.
Hofer, H. (1988). Variation in resource presence, utilization and reproductive success withina population of European Badgers (Meles meles).Mammal Rev.18: 25-36
Innes, H. S.(1978). Energetics of pelage polymorphism in the Eastern Gray squirrel (Sciuruscarolinensis). University of Guelph. PhD thesis.
Jackson, J.(1977). The annual diet of the Fallow deer (Dama dama) in the New Forest,Hampshire, as determined by rumen content analysis.J. Zool. , Lond.181: 465-473
Jackson, J.(1980). The annual diet of the Roe deer (Capreolus capreolus) in the New Forest,Hampshire, as determined by rumen content analysis.J. Zool.192: 71-83
212
Jackson, J., Chapman, D. I. & Dansie, O.(1977). A note of the food of Muntjac deer(Muntiacus reevesi). J. Zool. , Lond.183: 546-548
Johnson, I. P., Flowerdew, J. R. & Hare, R.(1992) Populations and diet of small rodentsand shrews in relation to pesticide usage. inPesticides, Cereal Farming and theEnvironment. The Boxworth Project(eds Greig-Smith, P., Frampton, G. & Hardy, T.)144-156 ( HMSO, London).
Jong, C. B., Gill, R. M. A., Wieren, S. E. v. & Burlton, F. W. E. (1995). Diet selection byRoe deerCapreolus capreolusin Kielder Forest in relation to plant cover.ForestEcology and Management79: 91-97
Jurajda, P., Prasek, V. & Roche, K.(1996). The autumnal diet of otter (Lutra lutra)inhabiting four streams in the Czech republic.Folia Zool.45: 9-16
Kemenes, I. & Nechay, G.(1990). The food of ottersLutra lutra in different habitats inHungary.Acta theriol.35: 17-24
Kikkawa, J. (1964). Movement, activity and distribution of the small rodents Clethrionomysglareolus and Apodemus sylvaticus in woodland.J. Anim. Ecol.33: 259-299
King, C. M. (1980). The weaselMustela nivalisand its prey in an English woodland.J.Anim. Ecol.49: 127-159
King, C. M. (1989)The Natural History of Weasels and Stoats( Christopher Helm Ltd,Bromley).
Knee, C. I. (1983). Squirrel energetics.Mammal Rev.13: 113-122
Kruuk, H. (1978). Foraging and spatial organisation of the European badger, Meles meles L.Behavioral Ecology And Sociobiology4: 75-89
Kruuk, H. (1978). Spatial organisation and territorial behaviour of the European badgerMeles meles.J. Zool.184: 1-19
Kruuk, H., Parish, T., Brown, C. A. J. & Carrera, J. (1979). The use of pasture by theEuropean badger (Meles meles).J. Appl. Ecol.16: 453-459
Lewandowski, K. & Nowakowski, J. J. (1993). Spatial distribution of brown hare Lepuseuropaeus populations in habitats of various types of agriculture.Acta theriol.38: 435-442
Loman, J. (1991). The small mammal fauna in an agricultural landscape in southern swedenwith special reference to the wood mouse Apodemus sylvaticus. Mammalia55: 91-96
213
MacDonald, D. W., Atkinson, R. P. D. & Blanchard, G. (1997). Spatial and temporalpatterns in the activity of european moles.Oecologia109: 88-97
MacKinnon, K. S. (1976). Home range, feeding ecology and social behaviour of the greysquirrel (Sciurus carolinensis, Gmelin). Oxford University. DPil thesis.
Marboutin, E. & Aebischer, N. J. (1996). Does harvesting arable crops influence thebehaviour of the European hare Lepus europaeus?Wild. Biol. 2: 83-91
Moller, H. (1983). Foods and foraging behaviour of Red (Sciurus vulgaris) and Grey(Sciurus carolinensis) squirrels.Mammal Rev.13: 81-98
Moore, N. P., Whiterow, A., Langton, S. D., et al.(1998). The Assessment of BadgerDamage to Agriculture and Forestry.Unpubl Report Project VC 03181-84
Moors, P. J. (1975). The food of weasels (Mustela nivalis) on farmland in northeastScotland.J. Zool. , Lond.177: 455-461
Moors, P. J. (1977). Studies of the metabolism, food consumption and assimilationefficiency of a small carnivore, the weasel (Mustela nivalisL.). Oecologia27: 185-202
Nagy, K. A. (1994) Field bioenergetics of mammals: what determines field metabolic rates?Aust. J. Zool.42: 43-53.
Neal, E.(1986)The natural history of badgers( Croom Helm, Beckenham).
Niethammer, J. (1978). Weitere Beobachtungen über syntope Wasserspitzmäuse der ArtenNeomys fodiensundN. Anomalus, Zeitschrift für Säugetierkunde - InternationalJournal of Mammalian Biology43: 313-321
Nolet, B. A. & Kruuk, H . (1994) Hunting yield and daily food intake of a lactating otter(Lutra lutra) in Shetland.J. Zool. Lond. 233: 326-331.
Overskaug, K., Knutsen, B. & Broseth, H.(1995). Habitat and area use by red fox Vulpesvulpes vixens in mid-Norway.Fauna Norvegica Series A16: 29-33
Pelz, H. J.(1989) Ecological aspects of damage to sugar beet seeds by Apodemus sylvaticus.in Mammals as Pests(ed Putman, R.J.) 34-48 ( Chapman and Hall, London).
Pepin, D.(1986). Spring density and daytime distribution of the european hare lepus-europaeus in relation to habitat in an open field agrosystem.Z. Saugetierkunde51: 79-86
Pernetta, J. C.(1976). Diets of the shrewsSorex araneusL. andSorex minutusL. inWytham grassland.J. Anim. Ecol.45: 899-912
214
Pernetta, J. C.(1976) Bioenergetics of British shrews in grassland.Acta theriol.33: 481-497.
Pigozzi, G.(1988). Diet of the European badger (Meles melesL.) in the Maremma NaturalPark, Central Italy.Mammal Rev.18: 73-75
Plesner-Jensen, S.(1993). Ecology and Behaviour of Small Mammals on Expanded FieldMargins.D. Phil. Thesis Univ. of Oxford38-64
Pollard, E. & Relton, J. (1970). Hedges, V. A study of small mammals in hedges andcultivated fields.J. Appl. Ecol.7: 549-557
Poulton, S. M. C.(1995) Small mammal populations in hedgerows: The relationship withseed and berry production. inField Margings: Integrating agriculture and conservation(ed Boatman, N.) 133-138 ( BCPC Monograph,
Pounds, C. J.(1981). Niche overlap in sympatric populations of stoats (Mustela erminea)and weasels (Mustela nivalis) in northeast Scotland. University of Aberdeen. PhDthesis.
Putman, R. J. (1986). Foraging by Roe deercapreolus-capreolusin agricultural areas andimpact on arable crops.J. Appl. Ecol.23: 91-99
Putman, R. J. (1986). Competition and coexistence in a Multi-species Grazing System.Actatheriol. 31: 271-291
Putman, R. J. (1986) Foraging by Roe deer in agricultural areas and impact on arable crops.J. Appl. Ecol.23: 91-99.
Putman, R. J. (1986)Grazing in temperate ecosystems: Large herbivores and the ecology ofthe New Forest( Croom Helm, Beckenham).
Putman, R. J., Culpin, S. & Thirgood, S. J.(1993). Dietary differences between male andfemale fallow deer in sympatry and in allopatry.J. Zool.229: 267-275
Putman, R. J. & Hemmings, G. J.(1986). Can dietary quality of free-ranging ungulates besimply determined from faecal chemistry?Acta theriol.31: 257-270
Putman, R. J., Pratt, R. M., Ekins, J. R. & Edwards, P. J.(1987). Food and feedingbehaviour of cattle and ponies in the New Forest, Hampshire.J. Appl. Ecol.24: 369-380
Reitz, F. & Leonard, Y. (1994). Characteristics of European hare Lepus europaeus use ofspace in a French agricultural region of intensive farming.Acta theriol.39: 143-157
215
Reynolds, J. C. & Aebischer, N. J.(1991). Comparison and quantification of carnivore dietby faecal analysis: a critique, with recommendations, based on a study of the FoxVulpes vulpes. Mammal Rev.21: 97-122
Reynolds, J. C. & Tapper, S.(1991). Roe deer and oilseed rape: the first winter.GameConservancy Review of 199098-102
Reynolds, J. C. & Tapper, S. C.(1995). The ecology of the red foxVulpes vulpesin relationto small game in rural southern England.Wild. Biol. 1: 105-119
Ricci, J. C. (1983). Suivi d'u lacher de liebres d'importation (Lepus europaeus Pallas) aumoyen de la radiotelemetrie: mortalite, dispersion et utilisatiion de l'espace.ActaOecologica. Oecol. Applic.4: 31-46
Richards, C. G. J., White, A. C., Hurrell, E. & Price, F. E. F. (1984). The food of theCommon dormouse,Muscardinus avellanarius, in South Devon.Mammal Rev.14: 19-28
Rogers, L.(1990). Small mammal diet and distribution in oilseed rape fields.AberdeenLetters in Ecology4: 12-13
Rogers, L. M. (1993). The ecology of small mammals in set-aside land.PhD Thesis,Aberdeen University
Rogers, L. M. & Gorman, M. L. (1995). The diet of the wood mouseApodemus sylvaticuson set-aside land.J. Zool. , Lond.235: 77-83
Rogers, L. M. & Gorman, M. L. (1995). The home-range size of wood mouseApodemussylvaticusliving in set-aside and surrounding semi-natural and crop land.J. Zool.237:675-678
Roper, T. J. (1994). The European BadgerMeles meles- Food Specialist or Generalist.J.Zool.234: 437-452
Roper, T. J., Findlay, S. R., Lups, P. & Shepherdson, D. J.(1995). Damage by badgersmeles-melesto wheattriticum-vulgareand barleyhordeum-sativumcrops.J. Appl.Ecol.32: 720-726
Roper, T. J. & Lups, P. (1995). Diet of Badgers (Meles meles) in Central Switzerland - Ananalysis of stomach contents.Zeitschrift für Säugetierkunde - International Journal ofMammalian Biology60: 9-19
Roper, T. J., Lups, P. & Lycett, S.(1989) Badgers as pests in English vineyards. inMammals as pests(ed Putman, R.J.) 207-208 ( Chapman and Hall, London).
216
Roper, T. J. & Mickevicius, E. (1995). Badger (Meles meles) diet - A review of literaturefrom the former Soviet-Union.Mammal Rev.25: 117-129
Rudge, M. R. (1967). The food of the common shrewSorex araneusL. (Insectivora:Soricidae) in Britain.J. Anim. Ecol.37: 565-581
Sheperdson, D. J., Roper, T. J. & Lups, P.(1990) Diet, food availability and foragingbehaviour of badgers (Meles meles, L.) in southern England.Zeitschrift fürSäugetierkunde - International Journal of Mammalian Biology55: 81-93.
Shore, R. F., Feber, R. E., Firbank, L. G., Fishwick, S. K., MacDonald, D. W. & Norum,U. (1997). The impacts of molluscicide pellets on spring and autumn populations ofwood mice apodemus sylvaticus.Agriculture, Ecosystems & Environment64: 211-217
Short, H. L. & Duke, W. B. (1971). Seasonal food consumption and body weights of captivetree squirrels.J. Wildl. Manage.35: 435-439
Skinner, C. A. & Skinner, P. J. (1988) Food of badgers (Meles meles) in an arable area ofEssex.J. Zool. , Lond.215: 360-362.
Southwood, L. L., Evans, D. L., Bryden, W. L. & Rose, R. J.(1993). Nutrient intake ofhorses in Thoroughbred and Standardbred stables.Australian Veterinary Journal70:164-168
Strachan, R. & Jefferies, D. J.(1993)The water vole Arvicola terrestrisin Britain 1989 -1990: its distribution and changing status ( The Vincent Wildlife Trust, London).
Sulkava, R.(1996). Diet of ottersLutra lutra in central Finland.Acta theriol.41: 395-408
Swift, S. M., Racey, P. A. & Avery, M. I. (1985) Feeding ecology ofPipistrellus pipistrellus(Chiroptera: Vespertilionidae) during pregnancy and lactation. II Diet.J. Anim. Ecol.54: 217-225.
Szmidt, A. (1975) Food preference of Roe deer in relation to principal species of forest treesand shrubs.Acta theriol.20: 255-266.
Taberlet, P. (1986) What barn owlTyto-albapellets can tell us about the ecology of smallmammals a case study in Bas Chablais Haute-Savoie France.Rev Ecol Terre Vie41(2-3):193-218.
Tapper, S. C.(1979). The effect of fluctuating vole numbers (Microtus agrestis) on apopulation of weasels (Mustela nivalis) on farmland.J. Anim. Ecol.48: 603-617
Tapper, S. C. & Barnes, R. F. W.(1986). Influence of farming practice on the ecology ofthe brown hare lepus- europaeus.J. Appl. Ecol.23: 39-52
217
Tarrant, K. A., Johnson, I. P., Flowerdew, J. R. & Greig-Smith, P. W. (1990). Effects ofpesticide applications on small mammals in arable fields, and the recovery of theirpopulations.BCPC Pests and Diseases173-182
Tattersall, F. H., MacDonald, D. W., Manley, W. J., Gates, S., Feber, R. & Hart, B. J.(1997). Small mammals on one-year set-aside.Acta theriol.42: 329-334
Taylor, J. C. (1969). Social structure and behaviour in a grey squirrel population. Universityof London. PhD thesis.
Tew, T. E. (1992). Radio-tracking arable-dwelling woodmice.Wildlife Telemetry(eds,Priede, G. & Swift, S.M.), 561-569 (Ellis Horwood, Chichester).
Tew, T. E. (1994) Farmland hedgerows: habitat, corridors or irrelevant? A small mammal'sperspective. inHedgerow Management and Nature Conservation(eds Watt, T.A. &Buckley, G.P.) 80-94 ( Wye College Press, Wye).
Tew, T. E. & MacDonald, D. W. (1993). The effects of harvest on arable wood miceapodemus-sylvaticus.Biological Conservation65: 279-283
Tew, T. E., MacDonald, D. W. & Rands, M. R. W. (1992). Herbicide application affectsmicrohabitat use by arable wood mice (apodemus-sylvaticus).J. Appl. Ecol.29: 532-539
Tew, T. E., Todd, I. A. & MacDonald, D. W. (1995) Field margins and small mammals. inField Margings: Integrating agriculture and conservation(ed Boatman, N.) 85-94 (BCPC Monograph 58)
Thirgood, S. J. (1995). The effects of sex, season and habitat availability on patterns ofhabitat use by fallow deer (Dama dama). J. Zool.235: 645-659
Tittensor, A. M. (1970). The red squirrel (Sciurus vulgarisL.) in relation to its foodresource. University of Edinburgh. Phd thesis.
Watt, J. (1995). Seasonal and area-related variations in the diet of ottersLutra lutra on Mull.J. Zool.237: 179-194
Watts, C. H. S.(1968). The foods eaten by wood mice (Apodemus sylvaticus) and bank voles(Clethrionomys glareolus) in Wytham Woods, Berkshire.J. Anim. Ecol.37: 25-41
Williams, O. B., Wells, T. C. E. & Wells, D. A. (1974). Grazing management ofWoodwalton Fen: seasonal changes in the diet of cattle and rabbits.J. Appl. Ecol.11:499-516
Wilson, C. J. (1993). Badger damage to growing oats and an assessment of electric fencingas a means of its reduction.J. Zool.231: 668-675
218
Wilson, G. Harris, S. & McLaren, G. (1997)Changes in the British badger population,1988 to 1997( People's Trust for Endangered Species, London).
Wolfe, A., Whelan, J. & Hayden, T. J. (1996). Dietary overlap between the Irish mountainhareLepus timidus hibernicusand the rabbitOryctolagus cuniculuson coastalgrassland.Biology and Environment96B: 89-95
Wolk, K. (1976). The winter food of the European water-shrew.Acta theriol.21: 117-129
Wolton, R. J. & Flowerdew, J. R. (1985). Spatial distribution and movements of woodmice, yellownecked mice and bank voles.Symp. zool. Soc. Lond.55: 249-275
Woods, R. D., Dunleavy, P. J. & Key, G. E.(1996). Small mammal activity in newhedgerows.Brighton Crop Protection Conference: Pests and diseases 1996445-446
Yalden, D. W. (1976). The food of the hedgehog in England.Acta theriol.21: 401-424
Ylonen, H., Altner, H.-J. & Stubbe, M. (1991). Seasonal dynamics of small mammals in anisolated woodlot and its agricultural surroundings.Ann Zool Fenn28: 7-14
Zubaid, A. & Gorman, M. L. (1991). The diet of wood miceApodemus sylvaticusliving ina sand dune habitat in north-east Scotland.J. Zool.225: 227-232
Zwahlen, R. (1975). Ein Beitrag zur Ernährungsökologie und zum Schadverhalten desEichhörnchens.Naturhistorisches Museum der Stadt Bern Jahrbuch5: 223-244
219
Appendix: Examples of results generated by Access queries
Information in these tables was generated from. Microsoft Access database queries. They arenot intended as an exhaustive tabulation of the data but rather as examples of the sort ofquestions that may be usefully asked of the database
Data on breeding cycle
Months when young are born.
No.Mnths
Start Fin Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
Badger 7 Jan Jul ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � � � � �Bank vole 7 Apr Oct � � � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � �Brown hare 9 Feb Oct � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � �Common dormouse5 May Sep � � � � ÿ� ÿ� ÿ� ÿ� ÿ� � � �Common Shrew 5 May Sep � � � � ÿ� ÿ� ÿ� ÿ� ÿ� � � �Fallow deer 3 Jun Aug � � � � � ÿ� ÿ� ÿ� � � � �Feral cattle 12 Jan Dec ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ�Feral goat 4 Jan Apr ÿ� ÿ� ÿ� ÿ� � � � � � � � �Feral sheep 3 Mar May � � ÿ� ÿ� ÿ� � � � � � � �Field vole 8 Mar Oct � � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � �Fox 3 Feb Apr � ÿ� ÿ� ÿ� � � � � � � � �Grey squirrel 6 Feb Jul � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � � � � �Harvest mouse 6 May Oct � � � � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � �Hedgehog 5 May Sep � � � � ÿ� ÿ� ÿ� ÿ� ÿ� � � �Horse 6 Apr Sep � � � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � � �Mink 1 May May � � � � ÿ� � � � � � � �Mole 4 Apr Jul � � � ÿ� ÿ� ÿ� ÿ� � � � � �Muntjac 12 Jan Dec ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ�Otter 12 Jan Dec ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ�Pipistrelle 2 Jun Jul � � � � � ÿ� ÿ� � � � � �Polecat 3 May Jul � � � � ÿ� ÿ� ÿ� � � � � �Pygmy shrew 7 Apr Oct � � � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � �Rabbit 8 Feb Sep � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � � �Red squirrel 7 Feb Aug � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � � � �Roe deer 3 May Jul � � � � ÿ� ÿ� ÿ� � � � � �Stoat 2 Apr May � � � ÿ� ÿ� � � � � � � �Water Shrew 5 May Sep � � � � ÿ� ÿ� ÿ� ÿ� ÿ� � � �Water vole 6 Apr Sep � � � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � � �Weasel 6 Apr Sep � � � ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� � � �Wood mouse 7 Apr Oct ÿ ÿ ÿ ÿ� ÿ� ÿ� ÿ� ÿ� ÿ� ÿ ÿ ÿ
220
Species givng birth earlyNo.Months
Start End
Badger 7 Jan JulBrown hare 9 Feb OctFeral cattle 12 Jan DecFeral goat 4 Jan AprFeral sheep 3 Mar MayField vole 8 Mar OctFox 3 Feb AprGrey squirrel 6 Feb JulMuntjac 12 Jan DecOtter 12 Jan DecRabbit 8 Feb SepRed squirrel 7 Feb Aug
Species giving birth lateNo.Months
Start End
Bank vole 7 Apr OctBrown hare 9 Feb OctCommondormouse
5 May Sep
Common Shrew 5 May SepField vole 8 Mar OctHarvest mouse 6 May OctHedgehog 5 May SepHorse 6 Apr SepPygmy Shrew 7 Apr OctRabbit 8 Feb SepWater Shrew 5 May SepWater vole 6 Apr SepWeasel 6 Apr SepWood mouse 7 Apr Oct
Species with a long breeding seasonMnths
Start End
Brown hare 9 Feb OctFeral cattle 12 Jan DecField vole 8 Mar OctMuntjac 12 Jan DecOtter 12 Jan DecRabbit 8 Feb Sep
221
Associations with habitat and crops
(A x means that the species is known to prefer the particular habitat type: the absence of ax doesnot mean that the habitat is avoided.)
cereals grassland roughgrazing
pasture woodland forestryplantations
riverhabitat
Badger x x xBank vole x x xBrown hare x x xCommon dormouse xCommon shrew x xFallow deer x x xFeral cattle x xFeral goat x xFeral sheep xField vole x x x xFox x x x xGrey squirrel x xHarvest mouse x xHedgehog x x xHorse x xMink xMole x x xMuntjac x xOtter xPipistrelle x x xPolecat x xPygmy shrew x x xRabbit x x x x x xRed squirrel xRoe deer x x xStoat xWater shrew xWater vole xWeasel xWood mouse x x
222
Body Weights1(alphabetical order)
Bodywt (g)
Lowrange
Highrange
Male LowMale
HighMale
Female LowFem
HighFem
Badger 10100 9100 11600 13900 6600 16700Bank vole 26.1 21.9Brown hare 3230 3430 2900 3500 3100 3750Common dormouse 18 15 26Common Shrew 8 5 14Fallow deer 67000 44000 46000 80000 35000 52000Feral cattle 300000 280000Feral goat 52400 41100 39000 65000 29000 52000Feral sheep 20100 25600 13800 19800Field vole 15 42 15 32Fox 6800 5700 5500 9300 4200 8200Grey squirrel 550 400 720Harvest mouse 7 6 8Hedgehog 1100 1600Horse 200000250000Mink 1150 650 840 1805 437 810Mole 110 85 87 128 72 106Muntjac 14800 12200 10500 18300 9000 15800Otter 10100 7000Pipistrelle 4.2 5.7 5.15 7.05Polecat 1111 689 800 1710 530 915Pygmy Shrew 4 2.4 6.1Rabbit 1500 1200 2000Red squirrel 300 220 440Roe deer 23900 22300 18000 28500 18000 28000Stoat 260 320 190 220Water Shrew 12 18Water vole 311 272 246 386 225 310Weasel 120 63 106 131 55 69Wood mouse 18 13 27
223
Body Weights2(ascending order of weight)
Bodywt (g)
Lowrange
Highrange
Male LowMale
HighMale
Female LowFem
HighFem
Pygmy Shrew 4 2.4 6.1Pipistrelle 4.2 5.7 5.15 7.05Harvest mouse 7 6 8Common Shrew 8 5 14Water Shrew 12 18Common dormouse 18 15 26Wood mouse 18 13 27Bank vole 26.1 21.9Field vole 15 42 15 32Weasel 120 63 106 131 55 69Mole 110 85 87 128 72 106Stoat 260 320 190 220Water vole 311 272 246 386 225 310Red squirrel 300 220 440Grey squirrel 550 400 720Mink 1150 650 840 1805 437 810Polecat 1111 689 800 1710 530 915Hedgehog 1100 1600Rabbit 1500 1200 2000Brown hare 3230 3430 2900 3500 3100 3750Fox 6800 5700 5500 9300 4200 8200Otter 10100 7000Badger 10100 9100 11600 13900 6600 16700Muntjac 14800 12200 10500 18300 9000 15800Feral sheep 20100 25600 13800 19800Roe deer 23900 22300 18000 28500 18000 28000Feral goat 52400 41100 39000 65000 29000 52000Fallow deer 67000 44000 46000 80000 35000 52000Horse 200000250000Feral cattle 300000 280000