Mammal Bible - Health and Safety Executive · 3 Introduction This compendium is intended as a companion volume to the earlier volume “Birds and farming” (Buxton and Crocker, 1996)

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.


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 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.


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




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.



( 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


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


Population:


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




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


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 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.


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.


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 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.



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


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.




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.




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.


%)

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



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


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.



( - 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



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.



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.


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


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



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.


%)

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




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


Otter Lutra lutra


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.



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.



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.


%)

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


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.



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


%)

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


Pygmy shrew Sorex minutus

4


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.


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.


%)

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




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


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.



( 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




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.




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


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.



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




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


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.



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


Water shrew Neomys fodiens


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.




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.


%)

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


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.



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


Weasel Mustela nivalis


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.



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


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.


Population: Stable


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



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


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


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


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


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



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



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


Seeds N= 66, % OCCURRENCE = 29%13

Moss N= 66, % OCCURRENCE = 24%11

Fungi N= 66, % OCCURRENCE = 23%10

58



Apr May Insects, larvae Hansson, 1971N= 66, % OCCURRENCE = 20%9


Lumbricids N= 66, % OCCURRENCE = 11%5


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


Seeds N= 87 % OCCURRENCE = 32%12

Lumbricids N= 87. % OCCURRENCE = 23%9


Insects, larvae N= 87, % OCCURRENCE = 17%7




Berries N= 87. % OCCURRENCE = 2%1

Sep Nov Seeds N= 108, % OCCURRENCE = 59%22


Fungi N= 108, % OCCURRENCE = ..37%14



Moss N= 108' % OCCURRENCE = 26%10

59



Sep Nov Grass Hansson, 1971N= 108, % OCCURRENCE = 17%6

Berries N= 108, % OCCURRENCE = 11%4


Graminoids N= 108, % OCCURRENCE = 2%1

Lumbricids N= 108, % OCCURRENCE = 2%1

Dec Feb Seeds N= 46, % OCCURRENCE = 61%25








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



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



May Jul Herbs Watts, 196815

Seeds 11

Fungi 7

Sep Nov Seeds 27

Herbs 25

Fruits mainly blackberry, elderberry, hawthornberry21

Fungi 11

62



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



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


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



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



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



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



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



Sep Pollen/spores Richards et al., 19845

Oct Nuts/berries 85

Leaves 10

Insects 5

68



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



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


Formicids 11

Geophilomorphs 11


Isopods 6

Coleopteranlarvae

6

May Lumbricids 72

Dipteran adults 62

70



May Isopods Churchfield, 198456

Gastropods 50

Chrysomelids 28

Acarines 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


71



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


Carabids 20


Araneids 13

Opilionids 13

Isopods 13

Geophilomorphs 7

Coleopteranlarvae

7

Sep Dipteran adults 95

Lumbricids 74

72



Sep Gastropods Churchfield, 198453

Isopods 37


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


Dipteran larvae 22

Acarines 22

Dec Dipteran adults 90

73



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



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



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


Slugs 12

Diptera larvae 12

Hemiptera 8

Opiliones 8

Lumbricidae 4

Coleoptera larvae 4

Hymenoptera 4

Araneae 4

Acari 4

Apr May Plant 28


Lumbricidae 14

Slugs 12

Hemiptera 8

Lepidoptera larvae 7

Diptera adults 5

Coleoptera larvae 3

Diptera larvae 2

76



Apr May Other insectlarvae

Rudge, 1968a2

Araneae 1

Opiliones 1

Jun Aug Plants 22



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


Hymenoptera 9

77



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



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



Jun Jul Diptera adults Rudge, 1968b2

Coleoptera 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


Chilopoda Lithobiidae 1%1

80



Sep Oct Araneae Rudge, 1968b1

Mar May Lumbricidae Rudge, 1968c% FREQUENCY in 327 gut contents. Devon, UK.22

Plant 16

Diptera larvae 12



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


81



Jun Aug Isopoda Rudge, 1968c7

Opiliones 6

Acari 5

Araneae 5

Slugs 4


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


Araneae 6

Slugs 6

Other larvae 5

Isopoda 3

82



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


Opiliones 5

Isopoda 5

Diptera larvae 5

Coleoptera adults Not identified.5

Coleoptera larvae 3

Acari 3

83



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



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



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



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



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



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



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


Quercus 0.7

Agrostis tenuis 0.6

Jun Other grasses 37

Agrostis tenuis 23

90



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



Oct Other grasses Putman & Hemmings,1986

16

Other shrubs 8.5

Agrostis tenuis 3.1

Ulex 2.9

Bracken & ferns 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


Rubus 0.9

Agrostis tenuis 0.6

92



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



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



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



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


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


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



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


18

Lychnis flos-cuculi 17.8


99



Jun Poa angustifolia &P. pratensis

Faber & Ma, 1986a6.7

Cytisus scoparius(flower)

Broom4.7


3

Animal material 2

Aug Poa angustifolia &P. pratensis

33


29

Holcus lanatus 12


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


17

Poa angustifolia &P. pratensis

14


100



Oct Undeterminedplant material

Faber & Ma, 1986a2

Seeds 0.7

Other 0.3

Dec Holcus lanatus 63



2.5

Other 0.7


0.3

Mar Deschampsiaflexuosa

Faber & Ma, 1986b% FRESH WEIGHT in stomach contents. Arnhem,Netherlands.

Wavy hair grass

92

Hypnumcypressiforme

2.9


2.2


1

Vaccinium myrtilla 0.8

Lignified stems 0.7

Jun Deschampsiaflexuosa

Wavy hair grass77

101



Jun Hypnumcypressiforme

Faber & Ma, 1986b7.3

Vaccinium myrtilla 6


4.4

Lignified stems 3.4


0.8

Animal 0.4

Aug Deschampsiaflexuosa

Wavy hair grass72

Hypnumcypressiforme

13

Vacciniummyrtillus berry

4.3

Lignified stems 4.2


3

Carex arenaria 1.2

Animal 1.1


1

Vaccinium myrtilla 1

Oct Deschampsiaflexuosa

Wavy hair grass78


11

102



Oct Undeterminedseed

Faber & Ma, 1986b3


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



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






Insects, adults N=69 %OCCURRENCE=6%2



104



Jun Aug Grass Hansson, 1971N=80 %OCCURRENCE=95%33







Fungi N=80 %OCCURRENCE=8%3


Sep Nov Grass N=193 %OCCURRENCE=100%44







Fungi N=193 %OCCURRENCE=9%4



Dec Feb Grass N=55 %OCCURRENCE=98%37





105



Dec Feb Seeds Hansson, 1971N=55 %OCCURRENCE=9%3



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


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


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


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


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


9.2

Insects 8.7

110


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



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



4.7

111


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


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



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



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



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



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



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



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



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



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



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



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



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



Jan Mosses Putman et al, 19878.5

Calluna vulgaris 6.7

Molinia caerulea 2.5


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


Mosses 13


Pteridiumapuilinum

2

Carex sp. 1

Molinia caerulea 1


Juncus sp. 0.5

125



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


Ulex sp. 10.2

Agrostis canina 6.2


Juncus sp. 1.2

Pteridiumapuilinum

0.8

Carex sp. 0.7


Other 0.5


Eriophorum sp. 0.3

Festuca rubra 0.2

Apr Other Gramineae 37


Mosses 15

Tree leaves 13

126



Apr Agrostis canina Putman et al, 19874.8


Juncus sp. 1.3

Ulex sp. 1.2


Carex sp. 0.8

Other 0.5

Festuca rubra 0.5

Pteridiumapuilinum

0.3


May Other Gramineae 54

Molinia caerulea 17

Agrostis curtisii 7.5


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



Jun Other Gramineae Putman et al, 198748

Molinia caerulea 22



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


Forbs 0.2

Jul Other Gramineae 50

Molinia caerulea 24



Pteridiumapuilinum

2.2

Eriophorum sp. 1.3

Other 1.2

Juncus sp. 0.8

128



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



Pteridiumapuilinum

6.5

Mosses 1.5

Other 1.5

Agrostis canina 1

Juncus sp. 1

Tree leaves 1


Festuca rubra 0.7

Carex sp. 0.7

Eriophorum sp. 0.5

Forbs 0.3

Sep Other Gramineae 45

129



Sep Agrostis curtisii Putman et al, 198724

Pteridiumapuilinum

9.7

Molinia caerulea 7


Agrostis canina 2.5

Mosses 2.2


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


Pteridiumapuilinum

6.5

Mosses 4.5

Tree leaves 3.3


Agrostis canina 2.7


130



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


Ulex sp. 3.3

Agrostis canina 2.7


Pteridiumapuilinum

1.5

Other 1.3

Eriophorum sp. 1.2


Festuca rubra 0.5

Carex sp. 0.3

Juncus sp. 0.2

Dec Other Gramineae 30

131



Dec Agrostis curtisii Putman et al, 198716

Tree leaves 13

Mosses 12


Ulex sp. 9

Pteridiumapuilinum

2.5

Agrostis canina 1.8



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


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


Mink Mustela vison

May Jul Molluscs Clode & Macdonald(1995)

4

Insects 4

134


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


Mole Talpa europea

Earthworms 50 185 HBMDaily energy requirements for 80 g mole.No seasonal data

136



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.


Fungi Occurred in 19% of samples.0.6

Conifers, deadneedles


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


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


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


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


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


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


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


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


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


Otter Lutra lutra

Jul Jun Birds Watt, 1995b0.6

Amphibians 0.5

Clingfish(Lepadogasterlepadogaster)

0.2

Conger eel(Conger conger)

0.2

146


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


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



Jun Aug Araneids Churchfield, 1994% FAECES containing items. (Totals may exceed 100.)Study of shrews inhabiting watecress beds. UK.

64


Isopods 48

Dipteran adults 40


Dec Feb Acarines 80

Isopods 60

Araneids 50


Collembolans 20

Coleopteranlarvae

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



Jan Mar Coleoptera larvae Pernetta, 197626

Opiliones 15

Aranae 7

Apr Jun Coleoptera adults 34

Aranae 24


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



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



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



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



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



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



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



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



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



Sep Nov Pine seed Zwahlen, 197516

Spruce buds 5

Dec Feb Spruce seed Winter.45

Pine seed 28

Spruce buds 16

159



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



Jan Festuca ovina Hearney & Jennings,1983

90

Agrostis tenuis 29


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



Feb Cynosuruscristatus

Hearney & Jennings,1983

32

Agrostis tenuis 31

Lolium perenne 18

Deschampsiaflexuosa

11


9


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


Cynosuruscristatus

17

162



Mar Lolium perenne Hearney & Jennings,1983

13

Deschampsiaflexuosa

9

Holcus lanatus 8


8

Phleum pratense 6

Other herbs 4


3

Quercus robur 3

Poa spp. 3

Rosa spp. 3


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



Apr Pseudotsugamenziesii

Hearney & Jennings,1983

5

Quercus robur 3

Phleum pratense 3

Other herbs 3

Crataegusmonogyna

3

Deschampsiaflexuosa

3

Holcus lanatus 3


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


5

Quercus robur 5


Deschampsiaflexuosa

3

164



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


Other herbs 4


2

Deschampsiaflexuosa

1

Juncus spp. 1


Lolium perenne 1

Jul Urtica dioica 100

165



Jul Rubus fruticosus Hearney & Jennings,1983

100

Festuca ovina 76

Agrostis tenuis 52

Betula spp. 8

Quercus robur 6

Other herbs 4



Poa spp. 3


2

Cynosuruscristatus

2


1


Crataegusmonogyna

1

Aug Urtica dioica 97

Rubus fruticosus 89

Agrostis tenuis 61

Festuca ovina 59

Betula spp. 7


166



Aug Holcus lanatus Hearney & Jennings,1983

4

Quercus robur 3

Other herbs 3


3

Rosa spp. 2

Poa spp. 2


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


3

Betula spp. 2

Crataegusmonogyna

1

167



Sep Rosa spp. Hearney & Jennings,1983

1


1


Juncus spp. 1

Oct Rubus fruticosus 99

Urtica dioica 87

Festuca ovina 60

Holcus lanatus 23

Agrostis tenuis 21

Poa spp. 7


7

Juncus spp. 3

Betula spp. 3


2

Quercus robur 2

Other herbs 1

Rosa spp. 1

Deschampsiaflexuosa

1

Nov Rubus fruticosus 84

Festuca ovina 71

168



Nov Urtica dioica Hearney & Jennings,1983

51

Holcus lanatus 21

Agrostis tenuis 18

Quercus robur 5


5

Poa spp. 3


3

Juncus spp. 2

Hedera helix 2

Rosa spp. 2

Lolium perenne 2


Other herbs 1

Crataegusmonogyna

1

Cynosuruscristatus

1

Dec Rubus fruticosus 99

Festuca ovina 83

Urtica dioica 36

Agrostis tenuis 25

Holcus lanatus 14

169



Dec Hedera helix Hearney & Jennings,1983

8


5


4

Quercus robur 3


Other herbs 3

Juncus spp. 2

Crataegusmonogyna

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



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


9

Scots pine 8

Ferns 6

Bilberry 3

Fungi 3

171



Sep Nov Other forbs Henry, 19781

Sitka spruce 1

Dec Feb Heather Winter.41

Scots pine 28


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



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



Apr Grasses Jackson, 198010

Ivy 7

Conifers 5

May Bramble/rose 35


Herbs 16

Grasses 7

Ivy 6

Heather 5

Conifers 1

Jun Bramble/rose 32

Herbs 16


Holly 14

Conifers 8

Grasses 8

Heather 4

Ivy 4

Jul Bramble/rose 40

Herbs 16


Other 10

Grasses 8

Heather 7

174



Jul Ivy Jackson, 19803

Holly 1

Aug Bramble/rose 40

Herbs 17


Other 10

Grasses 8

Heather 7

Ivy 3

Sep Bramble/rose 38

Herbs 17


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



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



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



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


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



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



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


Geophilomorphs 18

Isopods 16

Staphylinids 14

Gammarus 13

Ostracods 13

Araneids 13


Carabids 11

Acarines 11

Other dipteranlarvae

11

180



Mar May IndeterminateColeoptera adults

Churchfield, 198410.5

Trichopteranadults

5

Formicids 5

Lepidopteranlarvae

5

Simulium sp.larvae

5

Ephemeropteranlarvae

4.2


Hemipteran adults 2.6

Lithobiomorphs 1.3

Diplopods 1.3

Trichopteranlarvae (uncased)

1.3


1.3

Jun Aug Asellus 70

Dipteran adults 59

Gammarus 33


29


23

181



Jun Aug Other dipteranlarvae

Churchfield, 198419

IndeterminateColeoptera adults

19

Formicids 18



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



3.6

Tipulid larvae 2.9

182



Jun Aug Coleopteran adults Churchfield, 19842.9

Dermapterans 1.8

Lepidopteranlarvae

1.8

Sep Nov Asellus 63

Dipteran adults 56



40

Gammarus 31


29

Lumbricids 27

Gastropods 25


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



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


20


18


18

184



Dec Feb Araneids Churchfield, 198415


13

Acarines 8.6


Geophilomorphs 7.7

Tipulid larvae 7.7

Diplopods 6.3

Ostracods 5.4

Isopods 4.4

Ephemeropteranlarvae

4.3


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



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



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



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



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



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



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



Small rodents Day, 1968% FREQUENCY in gut. UK. (From Handbook of BritishMammals).

No seasonal data 55

Lagomorphs 18

Birds/eggs 16


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



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



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



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



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


196



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



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



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



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



Mar Mar Vegetative planttissue

Pelz, 198922

Sugar Beet seeds 7

Apr Apr Insect larvae Data from 49 individuals45

Earthworms 26


24

Cereal grain 5

May May Earthworms Data from 16 individuals40

Cereal grain 30


16

Insect larvae 10

Dicot seeds 4

Jun Jun Cereal grain Data from 15 individuals32

Dicot seeds 25

Insect larvae 25

Earthworms 9


9

Jul Jul Cereal grain Data from 10 individuals48

Insect larvae 28

Dicot seeds 16


8

201



Aug Aug Cereal grain Pelz, 1989Data from 41 individuals37

Insect larvae 28

Dicot seeds 20


10

Earthworms 5

Sep Sep Cereal grain Data from 18 individuals33

Insect larvae 25

Dicot seeds 20

Earthworms 13


9

Oct Oct Insect larvae 30

Cereal grain Data from 48 individuals30


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



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


2

Insects 1

Sep Nov Monocotyledons 50

203



Sep Nov Seeds Rogers & Gorman,1995

35

Other plantmaterial

6

Insects 5

Dicotyledons 2


1

Dec Feb Monocotyledons 68

Seeds 25

Insects 5


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



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



Sep Sycamore seed Watts, 196817

Flower 8

Other dicotyledonleaf

2

Oct Seed endosperm 79

Elderberry (fruit) 15


Bracken 1

Ash keys 1

Nov Seed endosperm 69

Sycamore seed 14

Fungi 7

Fibre fromsycamore keys

4

Hawthorn (fruit) 3


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 lepuseuropaeus.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

Documents

Mammal Bible - Health and Safety Executive · 3 Introduction This compendium is intended as a companion volume to the earlier volume “Birds and farming” (Buxton and Crocker, 1996)