The origins, and past and present status of the Little Owl

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THE ORIGINS, AND PAST AND PRESENT STATUS OF THE LITTLE OWL J. R. MARTIN The Little Owl (Athene noctua vidalli) is an introduced species to Britain. It is not normally present in Ireland, although it has been recorded there four times. The last occasion was in 1981. Breeding has never been recorded (Glue, in Gibbons et al., 1993). The Little Owl's natural ränge extends from the western coast of mainland Europe, across to central Russia, Mongolia and to China, although its ränge does not reach the Pacific Ocean. It is absent from India and much of south-east Asia. A paler race of the Little Owl, A.n.lilith, inhabits the dry, sandy region of the eastern end of the Mediterranean in Syria and Israel. The European race A.n.vidalli, is found in New Zealand where it was introduced by the Otago Acclimatisation Society in around 1906 (Marples, 1942). In India, parts of Indo-China and southern Iran, the Little Owl is replaced by the Spotted Little Owl (A.brama). The Forest Spotted Little Owl (A.blewitti) of northern India has not been recorded since 1968 and is endangered, perhaps even extinct, for despite searches in recent years it has not been recorded since 1968 (Burton, 1984).

Breeding Glue and Scott (1980) found that the Little Owl nests in a variety of places, but prefers to breed mainly in agricultural areas with plenty of timber in the form of copses, hedgerows, orchards or woods. They found that out of a total of 526 nests nearly 92% were in trees, and the most favoured site was a hole in a deciduous species. Buildings and other structures accounted for 6.7%, whilst rock clefts, rabbit burrows and other sites accounted for 1.7%. The data were obtained from the nest record cards held by the British Trust for Ornithology, and therefore represent the countrywide picture for nest sites and habitats in mainland Britain. R. B. Warren informs me that during the 1970s he found a nest in a quarry in east Suffolk. Jourdain (in Witherby et al., 1938) stated that the nests of Stock Dove (Columba oenas) and Jackdaw ( C o n us monedula) were also used. According to Glue and Scott (1980), and Wijnandts (1984) the Little Owl has the shortest breeding season of any of the five owls which regularly breed in Britain, with egg-laying almost exclusively confined to April and May. Glue and Scott (1980) found that 83% of clutches were started between 1 Ith April and lOth May, and Walpole-Bond (1938) gives dates in Sussex as between 20th April and lOth May. The size of the average clutch is 3.59, with an incubation period of around 28 to 33 days (Haverschmidt, 1946; Glue & Scott 1980). Fiedging occurs after 24 to 31 days, when young owls closely resemble their parents. Bearing these facts in mind, sightings of Little Owls in July may represent birds dispersing from the nest-site (Glue, 1973 and personal observations), and should not, therefore, be assigned as possible breeding records.

Diet Mikkola (1983) has observed "Fewer studies have been published

on the Little

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Owl's diet than on that of other European owls". However, he does point out the thorough investigation which was carried out in Britain by Alice HibbertWare (1938). Following Claims that the Little Owl was preying extensively on game and poultry birds, she carried out a study of the owl's diet using material collected by a wide and large network of fieldworkers. Some of the fieldworkers were recruited from Suffolk, and contributions were submitted from Barton Mills, Brandon, Ipswich, Lavenham, Mildenhall and Saxmundham. The investigation consisted of two parts. The first was undertaken during a fĂźll year, from February 1936 to 1937, whilst the follow-up took place from March to July 1937. Altogether 2,460 pellets were collected and examined; nest Contents and food larders were also submitted, along with 51 gizzards from specimens which were shot for the investigation. From her findings she was able to report that "The Little Owl feeds largely on insects at all times of the year, during all stages of growth and in all localities". However, her survey also revealed that the Little Owl eats small mammals and, occasionally, small birds. Cramp (1980) states that birds up to the size of thrushes are taken by the Little Owl, along with the occasional Moorhen (Gallinula chloropuys) and Magpie (Pica pica). R. B. Warren informs me that during Hibbert-Ware's survey he sent her a dead Jay (Garrulus glandarius) which he found at a Little Owl's nest. He does not know whether the Jay had been killed by one of the owls, or whether it had been collected as Carrion. The follow-up investigation, which took place in 1937, was directed specifically at predation by the Little Owl on game and poultry. The enquiry consisted largely of the examination of 26 gizzards from Little Owls shot on game estates. Nest and larder Contents, along with pellets, were also sent from four of the 1936 sites, and three new sites. In this second part of the enquiry laboratory work was intensive, and an extremely close search was made for traces of poultry and game-chicks. Hibbert-Ware's survey was comprehensive and well documented, and should have laid to rest any fears that poultry farmers and gamekeepers might have had about the impact of this introduced bird on their charges, for during the whole two year enquiry only one definite and one 'doubtful' gamechick, along with seven poultry chicks featured in the overall findings. However, as we shall see later, it took a long time for these facts to become accepted. In a review of Hibbert-Ware's survey, Ticehurst (1939) showed that beetles and other insects formed the bulk of the Little Owl's diet, with earwigs outnumbering all other species. And although it was found that birds were taken, this was chiefly in the spring and summer when young birds were plentiful, and the owls had young. The prey species consisted mainly of House Sparrows (Passer domesticus), Starlings (Sturnus vulgaris) and thrushes, including Blackbirds (Turdus merula). In addition to Hibbert-Ware's and Ticehurst's findings, there is evidence that an even wider spectrum of prey is taken. For example, on 14th June 1942, Burton (1943) saw a Little Owl rise from a verge near Walpole carrying "an enormous Frog", and Haverschmidt (1946) found that a pair he studied in Holland from 1938 to 1941 ate a number of frogs. On 16th June 1985, this

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PAST AND PRESENT STATUS OF THE LITTLE OWL

Linie Owl will: Earthworm.

Suffiilk. 194V.

(Photo Eric Hosking)

author observed a Little Owl catch and carry off to its nest a small Grass Snake (Natrix nalrix); and Buxton (1947) recorded a young headless Grass Snake about eighl inches long, at the entrance to a Little Owl's nest in Buckinghamshire. Haverschmidt also found that the birds he studied ate a number of earthworms, and this fact has also been reported by Hosking and Newberry (1945). In Britain earthworms do form a significant part of the Little Owl's diet, especially during the early stages of growth in the owlets [D. Glue pers. comm.]. There is no doubt though that larger prey is taken. In 1988 the author witnessed a Little Owl rising with considerable difficulty from a road, carrying a large rat in its talons. Whether the rat was Carrion, or whether it had been killed by the owl is not known. However, Hosking and Newberry have reported an instance where a young Little Owl was locked in combat with a rat. The owl was joined in the fight by one of its parents, and all three eventually disappeared into the owl's nest hole. Later, when the observer (Mr Phillips) looked into the nest he found "the rat there, dead, and covered with blood".

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Haverschmidt was also able to report on the first confirmed instance of cannabalism in the Little Owl, although as he states it had previously been suspected by Hibbert-Ware. History in Britain There is evidence that the Little Owl arrived in Britain prior to its known introduction in the 19th Century. For example, remains of the Little Owl have been recorded from inter-glacial deposits in Britain (Linn, 1979), and more recently Dr Eagle has written "In 1758 Edwards figured a Little Owl caught alive in a chimney near the Tower of London, and since that date until its successful introduction into England about a score of examples were recorded as having occurred in Kent, Sussex, Surrey, Middlesex, Norfolk, Cambs, Derbyshire, Yorks, Westmorland, Flint, Worcs, Wilts and Devon. Most if not all of these were doubtless immigrants from the Continent, it being certain that it does cross the North Sea occasionally, for one was captured on a smack 10 miles east of Yarmouth "s (Saunders, 1889). Turning to the 19th Century introductions, it was in May 1843 that Charles Waterton released five Little Owls in the grounds of his home at Waltham Hall in Yorkshire. However, it appears they did not survive (Lever, 1984). The first successful introduction was made at Stonewall Park, Edenbridge, in Kent, when several releases were made from 1874 onwards, but the first breeding record was not obtained until 1879. Further releases ensured that the Little Owl became eslablished south of the River Thames during the early 1900s. However, it was the releases by Lord Lilford, at Oundle in Northamptonshire, which really established the Little Owl as a breeding species throughout Britain. From 1888 he released large numbers of Little Owls in the vicinity of his home, until in 1895 he proudly boasted that he had succeeded in establishing the Little Owl as a resident breeding species in Northamptonshire. It seems likely that it was offspring from these owls which eventually colonised Suffolk. Migration Although the Little Owl is not credited with being a migratory bird there is evidence, as we have seen, that Little Owls have visited Britain from time to time, and there is evidence that this probably continues, for there can be no doubt that the Little Owl is capable of Aying long distances. For example, Hickling (1983) reported a British recovery from a flight of 182km, and Mikkola (1983) says that, although regulär migrations do not occur, some individual Little Owls do wander over distances of up to 300km. From ringing recoveries of 322 Little Owls of all age groups, Glue (in Lack, 1986) has calculated that 71% of Little Owls are found within 10km of their ringing place, and 2% are found 100km from their ringing place. Interestingly, Cramp (1990) makes no reference to Little Owl migration into Britain. In addition to these facts, and the historical records supplied by Babington (1886) and Ticehurst (1932), the Little Owl has been regularly recorded at Landguard, Felixstowe, since regulär monitoring has been in operation, and especially in the autumn (Table 1). Some or all of these records may, of course, relate to Suffolk bred birds, but it is possible that some are migrant birds and an

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PAST AND PRESENT STATUS OF THE LITTLE OWL

open mind must be kept until further evidence is available. The systematic ringing lists published in Suffolk Birds 1986-93, relating to owls ringed in Suffolk, show that only the Short-eared Owl (Asio flammeus) has received less attention than the Little Owl (Table 2). Clearly we need to find out more about the movements of this bird, and in 1992 a surprisingly high number of Little Owls were ringed. So, in time, this information may be forthcoming. Table 1: Little Owls recorded at Landguard, Felixstowe 1981-92 Oct/Nov 1981 1982 7th Sept-lOth Nov, four dates 30th Sept, 13th-15th Oct, 2nd and 18th Nov, Ist Dec 1984 6th Nov 1985 2nd, 41h, 17th Nov, 7th Dec 1986 1987 12th May, 21st Sept, 26th Oct 1988 At least two in October, at least one in November Ist April, 16th Oct 1989 8th-14th Oct, seven days in Nov, throughout Dec 1990 1991 Five dates in January, 20th Oct, 20th and 23rd Nov One, 22nd October 1992

1 singles singles 1 singles singles singles

The record in 1981 was the first since 1975. Additionally, a Single bird was flushed from the bushes at Minsmere sluice on December 19th, 1987. Source: Suffolk Birds Table 2: Owl species and numbers ringed in Suffolk 1986-1992 1986

1987

1988

1989

1990

4 1 7 13 1

6 1 4 2

7 1 3 5

5

-

-

Barn Owl Little Owl Tawny Owl Long-eared Owl Short-eared Owl Source: Suffolk Birds

1991

1992 -

1 2

4 1 3 3

-

1 3 4

8 4 6

-

-

-

-

-

Total 26 13 25 35 1

Recent history in Suffolk Lever (1984) stated that the first occurrence of the Little Owl in Suffolk was 1901, but clearly he was referring to descendants of the imported stock which Ticehurst (1932) listed. Ticehurst in fact reported that the first one was seen at Cavenham on 23rd April 1902. Others were subsequently recorded from Herringswell, December 1904, and Great Fakenham, February 1907. Lever did not mention the sightings before 1900 which Babington (1886) catalogued (Table 3), of which no doubt some relate to genuine vagrants. As has been stated, the Little Owl became established as a breeding bird in Britain after its introduction in the 19th Century. It is, however, reasonable to ask, since there were a number of records of this species occurring in Britain prior to this, why it did not naturally become established earlier. Presumably, like some other non-breeding bird visitors to this country, these immigrants were too few and too widely scattered to breed. Trans. Suffolk Nat. Soc. 30 (1994)


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Table 3: Little Owls reported in Suffolk prior to 1902 Date Not known cl840 February 1866 November 1874 cl877

Locality Yarmouth Holton Hengrave Thornham Oulton

No 1 alive 1 no data 1 female shot 1 1 no data

There are two other records of possible genuine vagrants reported by Babington. One was from Risby in 1866, although it may be that bird was a Tengmalm's Owl (Aegolius funereus). Another Little Owl was killed at Oulton on 26th October 1900. Regrettably there is little information in the literature which gives detail of the spread of the Little Owl across Suffolk. Ticehurst (1932) stated that the colonisation was exceedingly slow and gradual until the Great War. Between 1907 and 1912 the Little Owl had reached Sudbourne (1911) and Haiesworth (1912) where Ticehurst examined a nestling. DĂźring the Great War many gamekeepers were away in the armed forces and, due no doubt to the lack of persecution, its numbers increased. Payn (1978) stated that its population rose very markedly during the 1930s and early 1940s, when it was "certainly the commonest and most widespread owl in the County". Harvard (1931) considered the Little Owl to be "common anda nuisance", and there is no doubt that by then it was indeed common, even in East Suffolk. Edwards (1934) stated "! went to a little place on the north of Martlesham. The whole night long Nightjars and Little Owls kept up a discordant duet", and that same year Henniker (1934) reported that in the district of Eye there were "a great many Little Owls". This view of the Little Owl's status in Suffolk was not reflected throughout Britain however, for HibbertWare received several reports of Iocal declines in other counties during the course of her enquiry. However, I am reliably informed that the Little Owl was widespread in neighbouring Essex [R. B. Warren]. It has previously been mentioned that concern was expressed in Britain during the 1930s over the possible impact the Little Owl was having on gamebirds. At that time it was heavily persecuted, and debate began over whether the Little Owl should be protected. Even amongst naturalists the bird found little favour. For example, the committee which produced the report on raptorial birds in Suffolk stated "Little Owl. Is a destructive, and alien bird. Needs no protection." (Powell, 1935). Whether the Little Owl was the commonest and most widespread owl in Suffolk during the early 1940s is a matter for debate for many recorders were away in the armed forces and, as a result, it is probably that an accurate assessment would have been impossible. It is also likely that many Little Owls died of starvation during the severe winter of early 1940, and that further deaths occurred following the very severe winter of 1947. After the Second World War the debate intensified. At that time Iocal councils had the authority to give or remove protection to wildlife. For example, on 27th November 1946, it was reported from the Worthing Gazette that "By a considerable majority, it was decided that the little owl should he removed from the list ofwild birds deprived of protection" in West Sussex. This was a unique Trans. Suffolk Nat. Soc. 30 (1994)


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Situation in Britain, for as a report the previous day in the London Evening Standard stated "So from this week it is illegal to shoot the little owl or take its eggs in West Sussex. I know ofno other county council which have protected the little owl". In the following year it was reported by the Evening Standard on 6th March, that Kent County Council had "declared war" on the Little Owl, and had removed it from its protected species list. Later, on 2nd May, it was reported that the General Purposes Committee of the East Suffolk County Council had recommended that neither the "Little Owl nor its eggs be given protection at any period of the year", after "members had blamed the birdfor killing nightingales and newly-hatched chicks". This decision even reached distant readers, as this Stรถry was picked up by the Dundee Evening Telegraph and Post on 3rd May! It may be that East Suffolk County Council was influenced by a letter to the Eastern Daily Press, on 2nd May 1943, from R. H. B. Wilson, who, whilst remarking upon the decline of the Nightingale (Luscinia megarhyncho), wondered whether this was due to the increasing numbers of the Cuckoo (Cuculus canorus). He stated he was "duly informed by Mr Paterson (John Knowlittle) to 'watch the Little Owl'". He did, and considered it to be 'little wretch', and thought its importation was 'a great mistake'. Considering that the Nightingale frequents dense undergrowth in woods and copses, and the Little Owl is a bird of the open countryside, it seems difficult to imagine the Little Owl having any impact upon Nightingale populations. Indeed, later that year, on lOth July, the Evening Standard reported that "The little owl 'condemned' by East Suffolk County Council, is to be reprieved because it is 'a friend of the farmer'." It is clear that the law over the protection of owls in this country was in turmoil at that time, and it was not until the Protection of Birds Act, 1954, that all owls were fully protected. However, as Glue (1971; 1973) reported, persecution of the Little Owl continued, and clearly it was to take a while longer before the results of Alice Hibbert-Ware's study became firmly established. Apart from the debate over its protection in Suffolk and elsewhere, there is little in the literature which refers to the Little Owl during the 1940s; and in fact it was not until the first Suffolk Bird Report was published (Westall, 1952) that we have any idea of its status, when it was reported that it was a "Widespread resident", and "a further report of decline in numbers received." There is little additional information during the 1950s, although Payn (1952-3) reported from south-west Suffolk that it was a common resident. However, during the 1960s the Situation was to change significantly, and following the severe winter of 1962-3, the worst on record, the Little Owl declined sharply. For example, the year following that winter it was reported (Payn. 1964) that very few records had been received, and that numbers were steadily declining everywhere. Later Payn (1967) reported that it was "becoming steadily less plentiful", and this Situation continued, for it was further reported (Payn, 1968) that the Little Owl was "Now very scarce almost everywhere". This view continued throughout the remainder of that decade and most of the next. It was not until 1977 that it was "Recorded from

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54 parishes, a distinct improvement on past few years" (Payn, 1979). It seems likely that it took the Little Owl some while to recover from the very severe weather of the early 1960s, but it is also clear that modern farming methods were having an effect on its numbers. For example, Payn (1982) stated the Little Owl was "lost as a breeding species (at Härtest), because of the destruction of their nesting trees". It also seems very likely that the Little Owl suffered as a result of the application of highly toxic pesticides during that period. Being largely insectivorious it may well have suffered more losses as a result of poisoning than other owls. The loss of suitable nesting places undoubtedly helped in the Little Owl's decline. They often rely upon small farmland trees with narrow winding cavities, and pollard willows, especially along river-banks, to nest in. Many of these trees have been removed in recent years. Recent status of the Little Owl in Suffolk In the early 1980s the status of the Little Owl was not known. Moore (1981) reported that 59 records were submitted (twice the number of the year before), but wondered whether this reflected the fact that there were more birds, or simply more recorders. It was reported from 49 sites the following year (Moore, 1982), and breeding was noted at Minsmere, the first occurrence for 12 years. Since that time the number of reported sightings has gradually increased and Piotrowski (1993) reported it from 82 localities in 1991. More recently Sanford (1993) has shown that the Little Owl is widely but thinly distributed throughout Suffolk, and during the period of intense recording for the Suffolk Breeding Bird Survey, from 1987 to March 1993 (Fig. 1), it has been recorded in 43 of the 56 whole or part 10km Square (76.7%), with confirmed breeding in 31 (55.3%). Although the Little Owl is not as common as it was before the Second World War, it appears that it is a very much overlooked bird, and clearly there are many localities where there are Little Owls which have yet to be reported. Pairing According to Glue and Scott (1980) males do not Start to define their territories until the end of January or the beginning of February, with the main breeding activity beginning in March. Haverschmidt (1946) considered that a pair of Little Owls stay faithful to their territory throughout the year, and pair bonding, through copulation, occurs early in the year. He found it 'remarkable' that copulation could take place during December, even though egg-laying does not normally occur until the following spring. Through capturing and ringing individual Little Owls, over three and four year periods, both at the nest and using mist-nets, Glue and Scott considered that pair-bonding and site tenacity was strong, because this indicated that some individuals do occupy the same territory in successive years and throughout the year. They reported that certain sites had held Little Owls for over 25 years. From my experience a pair can often be located throughout autumn and early winter prior to breeding, when the cock bird arrives near to the nest-site

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PAST AND PRESENT STATUS OF THE LITTLE OWL

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and calls to the hen to join him. The pair then may 'sing' a vocal duet before Aying off together. In an area where a pair is observed during the winter period the site should be checked the following spring for signs of breeding. However, Glue and Scott stated that Little Owls mature early, and that it was not uncommon to find non-breeding and unmated individuals on territory throughout the breeding season.

Friend or foe? Lever (1984) applauded the introduction of the Little Owl into Britain stating that it is "unique among British vertebrates, in that it is the only orte of some sixty or so naturalised animals that is actually beneficial to man. As a largely insectivorous and mainly diurnal small bird ofprey, itfills an ecological niche left vacant by native raptors." When making these comments, Lever did not mention the impact the Little Owl may have on insect populations, nor whether it was competing for food with other animals. He did not refer to possible nest-site competition with other hole-nesting birds, nor did he take in to account predation by the Little Owl upon some of Britain's other wildlife. Sutcliffe (1990), for example, has proposed that the Little Owl is a major factor in the decline of the Storm Petrel (Hydrobates pelagicus) on Skomer Island. Using pellet analysis he was able to show the heavy predation of petrels by Little Owls. The Little Owl had previously been deported from Skokholm Island because of its impact upon the Storm Petrel population there. The question remains, what impact is the owl having on other species? Clearly these are areas which need to be investigated, for as Glue (1991) has rightly pointed out, the biology of the Little Owl poses many interesting questions, yet it has received scant attention. At the end of the Suffolk Breeding Bird Survey we may have a much better picture of its distribution in the County, but many questions will still remain to be answered.

Acknowledgements A number of people have been of great assistance to me. I am particularly grateful to Stanley Dumican, for his considerable help and encouragement, and to Colin Hawes and Marina Davies, for their comments. I am especially thankful to R. B. 'Bob' Warren, for telling me some of his experiences during the Little Owl Food Enquiry, and for making available to me Alice HibbertWare's correspondence to him. David Glue, of the British Trust for Orni_thology, has been most encouraging, and has made valuable and helpful comment on the draft. I sincerely thank him. I am also pleased that David Hosking has made available the photograph of the Little Owl which was taken in Suffolk by his late father, and for making available to me historical Information on the Little Owl which Eric had collected.

References Babington, C. (1886). The Birds of Suffolk. Proc. Suffolk Inst. Archaeology and Nat. History. Catling and Ranson, Bury St Edmunds. Burton, J. A. (ed.) (1984). Owls of the World. (2nd ed.) Peter Lowe, Wallingford.

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Burton, P. J. (1943). Little Owl's Food. Trans. Suffolk Nats. Soc. 5: 127. Buxton, E. J. M. (1947). Little Owl taking Grass-snake. Brit. Birds 40: 55. Cramp, S. (1985). The Birds of the Western Palearctic (Vol. 4). Oxford University Press. Edwards, P. (1934). The Voices of Dawn. Trans, Suffolk Nat. Soc. 2: 305. Gibbons, D. W„ Reid, J. B. and Chapman, R. A. (1993). The Atlas of Breeding Birds in Britain and Ireland: 1988-1991. Poyser, London. Glue, D. E. (1971). Ringing recovery circumstances of small birds of prey. Bird Study 18: 137-146. Glue, D. E. (1973). Seasonal mortality in four small birds of prey. Ornis. Scand. 4: 97-102. Glue, D. E. (1991). BTO Raptor and Owl Research Register. BTO News, 172: 5. Glue, D. E. and Scott, D. (1980). Breeding biology of the Little Owl. Brit. Birds 73: 167-180. Harvard, C. E. (1931). Bird Notes from a Buxhall Garden. Trans. Suffolk Nats. Soc. 1: 179-180. Haverschmidt, Fr. (1946). Observations on the breeding habits of the Little Owl. Ardea 34: 214-246. Henniker, Lord. (1934). Local Secretary for the Eye District's Report. Trans. Suffolk Nat. Soc. 2: 306. Hibbert-Ware, A. (1938). Report of the Little Owl Food Enquiry 1936-37. Witherby, London. Hickling, R. (ed.) (1983). Enjoying Ornithology. Poyser, Calton. Hosking, E. and Newberry, C. (1945). Birds of the Night. Collins, London. Lack, P. (ed.) (1986). The Atlas of Wintering Birds in Britain and Ireland. Poyser, Calton. Lever, C. (1984). The Little Owl in Britain. Hawk Trust Report 1984 14: 12-14. Linn, I. L. (ed.) (1979). Wildlife introductions to Great Britain. Report by the Working Group on Introductions of the UK Committee for International Nature Conservation. Nature Conservancy Council. Marples, B. J. (1942). A Study of the Little Owl, Athene noctua, in New Zealand. Trans. Royal Soc. New Zealand 72: 237-252. Mikkola, H. (1983). Öwls ofEurope. Poyser, Calton. Moore, D. R. (ed.) (1981). Systematic List. Suffolk Birds 1980. Suffolk Naturalists' Society, Ipswich. Moore, D. R. (ed.) (1982). Systematic List. Suffolk Birds 1982. Suffolk Naturalists' Society, Ipswich. Payn, W. H. (ed.) (1952-3). The Birds of South-West Suffolk. Trans. Suffolk Nat. Soc. 8: 82-100. Payn, W. H. (ed.) (1964). Systematic List. Suffolk Bird Report 1963. Trans. Suffolk Nat. Soc. 12: 490. Payn, W. H. (ed.) (1967). Systematic List, Suffolk Bird Report 1965. Trans. Suffolk Nat. Soc. 13: 242. Payn, W. H. (ed.) (1968). Systematic List, Suffolk Bird Report 1966. Trans. Suffolk Nat. Soc. 13: 431.

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Payn, W. H. (1978). The Birds of Suffolk (2nd ed.). Ancient House Press, Ipswich. Payn, W. H. (ed.) (1979). Systematic List, Suffolk Bird Report 1978. Trans. Suffolk Nat. Soc. 17: 306. Payn, W. H. (1982). The Breeding Birds of a Suffolk Garden 1941-1981. Suffolk Birds 1981. Suffolk Naturalists' Society, Ipswich. Piotrowski, S. H. (ed.) (1993). Systematic List. Suffolk Birds 1992. Suffolk Naturalists' Society, Ipswich. Powell, T. G. (1935). Report of the Raptorial Birds Committee. Trans. Suffolk Nats. Soc. 3: 69-71. Sanford, M. (ed.) (1993). Provisional Atlas of Breeding Birds in Suffolk. Suffolk Biological Records Centre, Ipswich. Saunders, H. (1889). An lllustrated Manual of British Birds. London. Sutcliffe, S. J. (1990). The Diet of Little Owls on Skomer. The Pembrokeshire Bird Report 1989. The Dyfed Wildlife Trust, Haverfordwest. Ticehurst, C. B. (1932). The Birds of Suffolk. Gurney and Jackson, London. Ticehurst, C. B. (1939). Hibbert-Ware on the Food on the Little Owl: a review Ibis III, 174-176. Walpole-Bond, (1938). A History of Sussex Birds. Witherby, London. Wijnandts, H. (1984). Ecological energetics of the Long-eared Owl Asio Otis. Ardea 72: 1-72. Witherby, H. F., Jourdain, F. C. R., Ticehurst, N. F. & Tucker, B. W. (1938). The Handbook of British Birds. Witherby, London. Westall, P. R. (ed.) (1952). Suffolk Bird Report 1951. Suffolk Naturalists' Society, Ipswich. J. R. Martin 17 Moss Way, West Bergholt, Colchester, Essex C 0 6 3LJ

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