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A PRELIMINARY SURVEY OF THE COLEOPTERA REDGRAVE A N D LOPHAM FENS R.

D.

OF

POPE

(British Museum [Natural History], London) and Lopham Fens have been acquired by the Suffolk Naturalists' Trust and are scheduled as a Site of Special Scientific Interest. It is intended that the area be managed by the Trust in such a way as to preserve the wild fauna and flora that it contains. In order to carry out this plan, many preliminary surveys of the area will have to be made and in May 1965, the British Museum (Natural History) was asked by the Lopham-Redgrave Fen Advisory Committee to investigate the beetle fauna. REDGRAVE

In view of the limitations imposed by justifiable resources, the survey was planned to cover one fĂźll season's collecting and part of another. A series of twelve, short visits was made, most of them comprising an hour or two's work on the day of arrival, two fĂźll day's collecting and a brief look round on the day of departure. The visits were regularly spaced from July to September, 1965, and from April to September, 1966. T h e number of collectors on each trip varied from three to four. T o work as many different types of habitat as possible while at the same time obtaining some idea of seasonal variations, certain areas were collected repeatedly, while part of each trip was devoted to covering new ground. In order to make the best use of the time available, as many collecting techniques as possible were used on each trip. Some places were worked with the sweepnet or beating tray, water collecting was by netting or panning. Vegetation around the edges of pools and along the river and drain banks was trodden into the water to discover its contents. Such vegetable debris as was found was either searched on the spot or samples were taken away for Berlese funnel extraction at the museum. Loose bark and dead timber were carefully examined, some reed-splitting tried and the few dung samples discovered were dissected in the field. Although some rabbit corpses and a few dead birds yielded their quota of beetles, by far the best results in this direction were obtained from the deliberate use of carrionbaited traps. Almost all the collecting was done in daylight, but one attempt at dusk produced some interesting results, including the only recorded buprestid. It was decided that an isolated list of captures would be of comparatively little interest. A faunal list is given below, but in addition the results are compared with those available for other similar areas and with the relevant county lists and records.


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T h e dates of capture for each species are given and a rough ecologi-

cal Separation is attempted. In this way it is hoped to give a picture of the beetle life of the fens throughout the season and in relation to the surrounding countryside.

The Fens The scheduled area (MAP 1) lies about five and a half miles west of Diss in Norfolk and runs along both banks of the river Waveney for about one and a half miles from its source. On the north bank are the Little, Middle, and Great Lopham Fens, on the south bank lies Redgrave Fen, Suffolk. They cover a little over 300 acres and include quite a variety of Vegetation (MAP 3). The reserve is largely unreclaimed peatland, but includes also fragments of the sandy plateaux on either side of the Waveney valley. Most of the area is in general flat, although in detail there are innumerable shallow pits and corresponding ridges on the surface. The surface of the fens proper has been lowered to an unknown extent by human agency and the microtopography is thus very largely man-made. Cross-valley drains, embankments, pits formed by peat cutting, and the wartime use of the fens as a bombing ränge provide their own special habitats. Some shallow ridges and islands of sand are important as they are acidic in contrast with the prevalent alkaline fen-peat. The vegetational communities may be divided into two groups, a freshwater series and a dry series. Freshwater series: the open aquatic habitat is represented by small pools, drainage ditches, and the river. In the wetter, badly drained areas, the dominant plants are either Cladium mariscus or Phragmites communis. Schoenus nigricans dominates the better drained parts of the fen proper. On the ridges between pools, Mollinia caerulea is dominant. On Redgrave Ridge and the margins of Little and Source Ridges (MAP 2) there is a mixed Community including plants typical of wet acid heath and rieh fen. Also present in these areas are species of Sphagnum, intermediate in pH requirements. Where disturbance of the land has occurred, such as has resulted from dredging Operations on the ditches, Epilobium hirsutum and Urtica dioica take over as dominant plants. Much sallow carr is developing in Middle and Great Fens, extending inward from the margins. In Redgrave Fen there is more mature carr.


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Dry series: just inside the northern boundaries of Little and Middle Fens there is a narrow strip of meadowland, some two acres of which has recently been cultivated. Other marginal areas support grasses, ling, bracken, broom, and gorse. Oak and birch scrub and woodland are to be found along the north and south borders of the reserve.

The Survey In the introduction, mention was made of the limited nature of the survey. It was essentially a sampling and skimming process. Little detailed ecological and biological information has thus been obtained, but a general foundation has been laid upon which more sophisticated structures may be erected at a later date. The records which follow present a complete list of the species captured during the survey and summarise all the available data appertaining to those captures. The names and their arrangement are approximately the same as used in "A Check List of British Insects" by Kloet and Hincks (1945). Family groups differ occasionally in accordance with more modern opinion. The Staphylinidae are listed in accordance with Tottenham (1949) and such nomenclatural changes as have been published since 1945 in other groups have been followed. Species within genera are arranged alphabetically in order to assist non-specialists. In order to relate the captures to the fauna of the surrounding countryside and that of similar wetland areas, cross-checks have been made against published lists and later supplements concerning the coleoptera of Wicken Fen, Cambridgeshire, Wood Walton Fen, Huntingdonshire, Oulton Broad, near Lowestoft, Suffolk and the various county records available for Norfolk and Suffolk. Precise references to these sources are given in the bibliography (see page 40). T h e Lopham-Redgrave species are here broadly divided into those more often found in damp situations than otherwise and "others". The lists of species found at Wicken, Wood Walton, and Oulton Broad have been analysed in the same way. It is of interest to see how the results of these analyses (TABLE on page 31) compare with the vegetational pattern of each area.


COLEOPTERA OF REDGRAVE AND L O P H A M

A

C O M P A R I S O N O F L O P H A M - R E D G R A V E F E N S W I T H OTHER E A S T F E N S AND W I T H T H E I N C L U D I N G C O U N T I E S

From Lopham-Redgrave Fens Common to Lopham-Redgrave, Wicken and Wood Walton Fens In common with Wood Walton Fen In common with Wicken Fen In common with Oulton Broad Not recorded from Wood Walton Fen Not recorded from Wicken Fen Not recorded from either Wicken or Wood Walton Fens Species new to Norfolk Species new to Suffolk Species new to both counties

31

FENS

ANGLIAN

Number of Species Recorded Total Wetland "Others" 771 226 545 334

140

194

518 509 180

152 192 57

366 317 123

253 262

75 33

178 229

178 113 96 46

22 — — —

156 — — —

The places and dates of capture for each species are set out in the appropriate columns of the main species list. On the extreme right of the list there is for each species a coded summary of the bionomic data recorded at the times of its capture. This gives some idea of the adult life-span and its ränge of habitat insofar as Lopham-Redgrave Fens and the seasons 1965 and 1966 are concerned. Certain incongruities appear among the habitat block records. These are species obviously out of their normal environment. T h e records are kept in for completeness, but noted as accidents by means of an exclamation mark (!). The Table given above summarises the results of analyses made from the records set out in the main species list. Discussion Lopham and Redgrave Fens occupy a comparatively small area, being about half the size of Wicken Fen and about three-fifths as large as Wood Walton Fen. As can be seen from the map on page 29, quite a large part of the 300 or so acres is already too dried out to be called true fen. T A B L E SUMMARISING ECOLOGICAL ANALYSES

Approx. Acreage Lopham- Redgrave Fens Wicken Fen Wood Walton Fen Oulton Broad

314 615 514 100+

Species Recorded

Families Recorded

Wetland Species

"Others"

771 1,069 728 438

61 58 51 38

226(29%) 415(39-6%) 263(36%) 120(27-4%)

545 654 465 318

Apart from a single record (Apion simile Kirby) for Redgrave Fen mentioned in Fowler and a small number of species taken by Dr. A. Eve in 1957, the area appears to have been completely neglected by coleopterists. By contrast, Wicken Fen has been a happy hunting ground for entomologists since early in the nineteenth Century and the first records for Wood Walton Fen date


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from 1913. In both Wicken and Wood Walton there are mown rides and access to ponds and ditches is comparatively simple. Also the heaps of Vegetation produced by mowing and cutting provide excellent habitats for a large number of species, acting as a concentrator in favour of the collector. There has been no recent mowing or cutting at Lopham-Redgrave. T h e only activity which has opened the area to any extent has been dredging and reed-cutting in the river by the East Suffolk and Norfolk River Board. In consequence the terrain is more difficult and the fauna more thinly distributed. Bearing this in mind, the total n u m b e r of species taken would seem to show that a fairly thorough search was made du ring the period under discussion. T h e justification for a series of short visits, involving a wealth of material (15 to 20,000 specimens) for sorting, mounting, and preparing for identification, is that no trip failed to add new species to the records and that each visit produced at least ten species which were never found again during the survey. In the comparison table on page 31, the most significant entries are those which demonstrate a lack of faunal overlap between Lopham-Redgrave and the other selected areas. Buck and Gardner (1962) remark on the 200 or so species that have been recorded from Wood Walton, but not from Wicken Fen. In the present survey the most important result is the recording of twenty-two species of wetland characteristic not yet found at either Wicken or Wood Walton. T h e following notes concerning these particular captures may be of interest. DYTISCIDAE

Hydroporus nigritus (F.). Three examples taken, two from small pools, one from the river. This species is said to be confusable with H. discretus and H. pubescens. It is hardly likely that this is the reason for its absence from the Wicken and Wood Walton lists. It is a widespread species in this country and was recorded from Quy Fen, Cambridgeshire by Power. F. BalfourBrowne (1926) included the species in his account of the aquatic Coleoptera of the "Wicken Fen area", but it does not appear in the list given by J. Omer-Cooper and C. E. Tottenham (1932). It is assumed that Balfour-Browne did not take the species in the fen itself and that the later collectors failed to find it there also. Hydroporus tieglectus Schaum. Four examples taken, one of them apparently in the sweepnet! This rather rare species is recorded by Balfour-Browne (1940) as occasionally common in east Norfolk. It does not appear in the county records for Cambridgeshire and Huntingdonshire. Ilybius subaeneus Er. A single example of this rarely recorded species was taken from the river Waveney. First recorded from


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33

Norfolk and SufFolk in the late nineteenth Century. Taken by Balfour-Browne in 1904 from Burwell Fen and in 1918 and 1925 from Quy and Bottisham Fens (Balfour-Browne 1950). HYDROPHILIDAE

Laccobius atrocephalus Reitter. A single specimen dredged from the river in early August. Apart from Cambridge, this is the only recorded locality for the species in eastern England. Helophorus nubilus (F). A single example, taken apparently in the sweepnet. Balfour-Browne (1958) records that he has taken the species on nineteen occasions since 1903. Helophorus walkeri Sharp. Two examples. Helophorus strigifrons Thomson. One example. These two records are inserted as the result of separate identications, kindly undertaken for the author by Mr. R. B. Angus. Their absence from the Wicken and Wood Walton lists is explained by their being unrecognised as distinct species when the records were compiled. As yet there is insufficient available data to assess the British distribution of either one. SCYDMAENIDAE

Neuraphes angulatus Müller. A single example discovered by Berlese funnel extraction of a sample of very wet vegetable debris from Great Fen. Although unrecorded from Huntingdonshire, N. angulatus has been taken on two occasions in the Cambridge area by Nicholson. Neuraphes elongatulus Müller. A single example from wet Vegetation on the bank of the river Waveney. T h e species is not recorded from either Cambridgeshire or Huntingdonshire. STAPHYLINIDAE

Micropeplus tesserula Curtis. A single example from under the bark of a fallen willow tree. Recorded from "the Fen districts of Cambridgeshire" by Fowler. Not recorded from Huntingdonshire. Tachyporus tratisversalis Grav. Taken on three occasions in quite different parts of the fens. Surprisingly absent from the Wicken and Wood Walton lists. Fowler does not imply that there is anything unusual in the distribution of this species. Being of distinctive appearance, it is not likely to have been confused with any other British member of the genus. Atheta (Stethusa) aquatica Thomson. A single example from rotting Vegetation very near the road entrance on the north boundary of Middle Fen. Recorded from both Cambridgeshire and Huntingdonshire. Fowler gives quite a wide ränge of


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localities for this species. As with other species of Atheta, the lack of records may simply reflect the extreme difficulty of naming many of them. Atheta (Brundina) gemina Er. This apparently rare species was taken by sweeping herbage along the drainage ditch between Middle and Great Fens. It is recorded from Cambridgeshire, but not from Huntingdonshire. Calodera aethiops (Grav.). Three examples, two by sweeping, one from leafmould in woodland. Recorded from Quy Fen, Cambridgeshire, but not from Huntingdonshire. Fowler regarded the species as local, but fairly widespread. PSELAPHIDAE

Reichenbachia impressa (Panzer). Two examples, one by sweeping, one by searching at the roots of Cladium. The species is unrecorded for both Cambridgeshire and Huntingdonshire. A widespread species, found in haystack refuse as well as in damp vegetable matter at the edges of ponds, etc. PTXLIIDAE

Ptenidium fuscicorne Er. Many of the Ptiliidae taken at the fens appear to belong to this species. However, the identification of any members of the group is at present little more than tentative and so biological attributes are also doubtfully accurate. It is perhaps a little optimistic to add P. fuscicorne to the list of wetland species collected at Lopham and Redgrave Fens. HELODIDAE

Cyphon hilaris Nyholm. This species was added to the British list by Kevan in 1962. It was previously recorded as C. ochraceus Stephens, with which it was confused. It is very common at Lopham-Redgrave and is doubtless equally plentiful at the other two East Anglian fens. CRYPTOPHAGIDAE

Telmatophilus schoenherri Gyll. A single specimen, taken in the sweepnet, and one other discovered by Splitting stems of Typha. There appear to be no records of this species from Huntingdonshire, but Fryer took it at Chatteris and Horsway in Cambridgeshire. CHRYSOMELIDAE

Galerucella nymphaeae (L.). A single example from the wettest part of Great Fen. This species has been recorded several times from Cambridgeshire, although not from Wicken Fen. There are no records of the species from Huntingdonshire. Fowler regarded the species as local, but fairly widespread.


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35

CURCULIONIDAE

Acalyptus carpini (F.). A single example beaten from scrub in Great Fen. Taken on Burwell Fen by Power, unrecorded from Huntingdonshire. Fowler considered the species as rare and gave only three localities. The G. C. Champion collection includes a series from near Oxford and Donisthorpe took the species at Weston-on-the-Green, Oxfordshire. Mr. P. M. Hammond, Recorder of Coleoptera for Essex, has told the writer that he took three specimens at Wicken Fen on 29th May, 1961. Amalorrhynchus melanarius (Stephens). Taken on two occasions, one specimen actually from the river! There are no records from Huntingdonshire. Nicholson took the species at Fen Ditton, Cambridgeshire. Gymnetron villosulus Gyll. One specimen taken by sweeping herbage along the river bank. Recorded from Cambridgeshire, but not from Huntingdonshire. In Fowler's day the species was regarded as very local. Phytobius quadricornis (Gyll.). Three examples of this rare species were taken, all near the north-west corner of Great Fen. Within the carr, not far from the place of capture, there is a very badly drained area supporting many aquatic plants. It is most probable that the species has its habitat there. It does not figure in the records of either Cambridgeshire or Huntingdonshire. Fowler gives London (Stephens), Sheerness, Kent, and Broxbourne, Herts, as the only localities known to him. The Donisthorpe collection includes a series from Rickmansworth, Herts. The Champion collection houses a series from Newbury, Berks, and a single example labelled "Norwich ex coli. R. Brown". If this last is correct, it represents a hitherto unpublished record from Norfolk. As the Norfolk-Suffolk boundary runs through the fens, the only county additions regarded as noteworthy are the forty-nine hitherto unrecorded from both. As has been stated earlier, the county records are set up on the basis of the lists published in the Victoria County Histories and supplements in the Journals of the Norfolk and Suffolk Natural History Societies. It is thus quite possible that individual records have been overlooked and that the number of new records resulting from this survey is in fact considerably smaller. The ecological analyses of the wetland areas compared shows Lopham-Redgrave to have a more varied fauna than either Wicken or Wood Walton Fens. More families have been recorded and the percentage of typically wetland species is lowest. There may be various explanations for this. The collecting may have been biased in favour of the easier parts of the terrain. The chosen


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seasons may have been somewhat abnormal. The watertable in Lopham-Redgrave Fens shows a very marked seasonal Variation. At the height of summer the river may be three or four feet below the general level of the fens. In winter, a great deal of Little Fen and parts of Middle and Great Fens are submerged. This will undoubtedly affect the fauna. As has been mentioned earlier, no mowing or cutting is carried out now. Many species inhabit heaps of cut and decaying Vegetation, whereas an area of Standing sedge supports a very meagre beetle population. A breakdown of the records showing how the larger families are represented in each of the three fen areas gives the following results: Lopham-Redgrave Carabidae 63 Dytiscidae 47 Hydrophilidae 38 Staphylinidae 200 Chrysomelidae 61 Curculionidae 111 (including Attelabidae and Apionidae)

Wood Walton

Wicken

90 60 44 148 69 107

136 67 54 275 85 117

The relatively small number of carabid species recorded from Lopham-Redgrave may be a reflection of the particular seasons, but it is more likely to be connected with the absence of litter heaps where so many of them have been taken at Wicken and Wood Walton. Only a third of the species are wetland, compared with 44% of those on the Wicken list and 43% of the total species from Wood Walton. A comparatively limited time was devoted to collecting from the open aquatic habitats offered by the fens. This may be the reason for the Lopham-Redgrave complex appearing at the bottom of the list, but it is also quite possible that river pollution, from saline effluent pumped in near the source from a watersoftening plant, has depleted the fauna. Also, a large number of small pools were never visited, some for lack of time, some from ignorance of their existence until it was too late. It would seem very probable, as both Norfolk and Suffolk support some 130 species of aquatic Coleoptera, that further collecting at LophamRedgrave will add a number of species to the present list. If any of the pools are cleared of surrounding Vegetation and so exposed to light and air, it is almost certain that the dytiscid populations will increase. The Staphylinidae collected at Lopham-Redgrave separate into a surprisingly large number of species. The proportion of wetland species is 25% against 33% of the Wicken list and 27% of all those recorded from Wood Walton. Many of the species were collected from rather dry situations, or from specialised habitats such as fungi or Carrion. Curiously, the number of species of Stenus


COLEOPTERA OF REDGRAVE AND LOPHAM FENS

37

is exactly the same in both Lopham-Redgrave and Wood Walton. The latter list includes five wetland species not recorded from Lopham-Redgrave. Numerically, this is matched by five wetland species from Lopham-Redgrave which are as yet not known from Wood Walton. Of the 116 species of Wicken Staphylinidae not found at either of the other two areas, thirty-eight are characteristic of wetland. Although the numbers of recorded species of Curculionidae from the three fen areas are very close, only thirty-nine are held in common. Lopham-Redgrave supports thirty-three species not yet discovered at either Wicken or Wood Walton. Of these, only four are wetland species. They have been discussed on page 35. Taken as a whole, the Lopham-Redgrave Curculionidae include 14% wetland species as compared with 22% of the Wicken list and 17% of the Wood Walton records. In the case of the Chrysomelidae the picture is different again. The Wicken list is markedly larger than that for either LophamRedgrave or Wood Walton. It includes a dozen species of Donaciinae against two from Lopham-Redgrave and none at all from Wood Walton. This raises the percentage of wetland chrysomelids to forty-seven for Wicken, whereas the ecological analyses for Lopham-Redgrave and Wood Walton give 18% and 20% respectively. Of the Lopham-Redgrave list, only twentyfour species are also found at Wicken and Wood Walton. Of the twelve species unique to Lopham-Redgrave, only one is a wetland species. This is Galerucella nymphaeae (L.) and has already been discussed on page 34. Buck (1962) suggests that the absence of Donacia and Plateumaris from Wood Walton may b e due to the regulär C l e a r i n g of reed from the drains. In Lopham-Redgrave this may well be part of the story, but equally the watertable fluctuations may render the area unsuitable for the satisfactory development of the larvae. Summary The first attempt at a survey of the Coleoptera of LophamRedgrave Fens has produced a species total of 771. This is the result of about thirty days' collecting by three or four people. A detailed examination of the species list and a comparison with the species lists of other East Anglian fens has shown the area to have a markedly varied fauna with a rather small percentage of species characteristic of wetland. This accords well with the fact that the scheduled site includes land which ceased to be fen many years ago and a considerable portion of the remainder is already fen in decline. T h e cessation of reed-cutting and the dredging of the river and ditches have also played their part in the evolution of the area.


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Acknowledgements The Redgrave Fen Advisory Committee has been mentioned in the introduction. The author's thanks are also due to the Earl of Cranbrook who initiated the project and encouraged its execution; to the Director and Trustees of the British Museum (Natural History) and the Keeper of Entomology in that institution, who both encouraged the enterprise and gave the author all the administrative assistance for which he asked. Seven members of the museum staff, namely R. G. Adams, M. E. Bacchus, M. J. D. Brendell, J. J. Brightman, M. Ford, B. Levey, and Miss E. R. Tozer, all took part in the various collecting trips and bore at least their share of the long task of preparing and identifying the material collected. Without their Willing Cooperation, the survey would have been impossible. The long list of Atheta species recorded is the result of Mr. H. Last's labours with this most difficult group for which the author is deeply grateful. Two Helophorus records, as acknowledged earlier, are identifications kindly supplied by Mr. R. B. Angus. The author is also indebted to the Management Committee of Lopham-Redgrave Fens for permission to make use of the text of the Management Plan in preparing this paper and to reproduce the maps on pages 27, 28 and 29. The collecting and subsequent work of the survey has been spread over many months and a special mention must be made of the tolerance and forbearance of the author's colleagues during the inevitable dislocation of the regulär work undertaken by the Coleoptera section in this museum. The material collected during the course of the survey is at present housed in the British Museum (Natural History), London. It is intended that a reference collection be stored at Fiatford Mill Field Station, Fast Bergholt. The complete field data associated with the survey is also stored at the British Museum (Natural History).

Key to Abbreviations used in the Species List Chart T O W Ww B

Characteristic of a wetland environment. Not classifiable as " T " above. Recorded from Wicken Fen. Recorded from Wood Walton Fen. Recorded from Oulton Broad.


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N S L M G R

39

Recorded f r o m Norfolk. Recorded f r o m Suffolk. Little Fen, L o p h a m . M i d d l e Fen, L o p h a m . G r e a t Fen, L o p h a m . Redgrave F e n .

Numbers 1 to 12 refer to individual collecting trips. corresponding dates are:—

The

(1) 7-9.vii.1965. (2) ll-13.viii.1965. (3) 8-10.ix.1965. (4) 20.iv. 1966. (5) 10-13.V.1966. (6) 23-25.v.1966. (7) 7-10.vi.1966. (8)28.vi.l.vii.1966. (9) 20.vii.1966. (10) 2-4.viii.1966. (11) 16-18.viii.1966. (12) 13-15.ix.1966.

Habitat Block Records The unbracketed numbers under this heading on the List of Species refer to the information given below. The numbers in brackets are the occasions upon which the species was taken under the circumstances referred to by the adjacent unbracketed number. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30.

Living wood. W o o d l a n d , scrub, isolated trees and bushes. H e r b a g e in general, not on acid heath areas. Usually by sweeping. H e r b a g e o n acid h e a t h areas, heather, ling, etc. Usually by sweeping. T a k e n f r o m individual flowers, usually u m b e l s . D e a d timber, Standing and fallen. T h i s includes hibernating specimens. U n d e r bark. I n c l u d i n g hibernating specimens. Associated with fungi, both mycelia and fruiting bodies. F r o m felled oak b r a n c h e s with t h e leaves still attached. F r o m birds' nests. F r o m building r u b b l e in woodland, n o r t h m a r g i n of M i d d l e F e n . F r o m u n d e r stones in arable areas. F r o m roots of grass. F r o m moss. F r o m plant roots and bryophytes in acid heath area. F r o m d a m p vegetable debris. F r o m very wet vegetable debris and Vegetation. (Around pools, etc.) F r o m stems of Typha. F r o m m u d d y banks of drains and river. F r o m waterweed dredged f r o m river. D r e d g e d f r o m small pools. F r o m drainage ditches. F r o m river—slow moving water. F r o m weir (see MAP)—fast moving water. F r o m a heap of rotting potatoes in M i d d l e F e n . F r o m d u n g . H o r s e droppings in Middle Fen, m e a d o w area. F r o m carrion-baited traps. Baited with rabbits or rats. F r o m naturally occurring Carrion. Rabbits, chicken, m o o r h e n , duck, mole, pigeon. F r o m t h e m o u t h s of rabbit burrows. F r o m " L o n g w o r t h " traps set to catch m a m m a l s . By sweeping and beating Vegetation at dusk.


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Bibliography Balfour-Browne, F. (1926). T h e Aquatic Coleoptera of the Wicken Fen area, Cambs. The Natural History of Wicken Fen part 3, 201-214. Balfour-Browne, F. (1940, 1950, 1958). British Water Beetles vols. 1, 2, and 3. Ray Society Monographs Numbers 127, 134, 141. Bedwell E. C. (1899). Coleoptera at Oulton Broad and District. Entomologist's Ree. J. Var. 11, 298-300, 335-38. Buck F D (1961). A Provisional List of the Coleoptera in Wood Walton Fen, Hunts. Proc. S. Lond. Ent. Nat. Hist. Soc. 1962, 93-117. Donisthorpe, H. J. K. (1938). The Victoria History of the County of Cambridgeshire and the Isle of Ely 1, 104-137. Edwards, J. (1901). The Victoria History of the County of Norfolk 1, 110-135. Elliott, E. A. (1936). Critical Notes on our Beetles. Tram. Suffolk Nat. Soc. Framlingham 3, 121-128. Fowler, W. W. (1887-1913). The Coleoptera of the British Isles vols. 1-5 and (suppl.) 6. London. Hammond, P. M . (1964). T h e Beetle Lathridius bifasciatus Reitter in Essex. The Essex Naturalist 31, 172-175. Kevan, D. K. (1962). T h e British Species of the genus Cyphon Paykull (Col. Helodidae), including three new to the British List. Ent. mon. Mag. 98, 114-121. Kloet, G. S. and Hincks, W. D. (1945). A Check List of British Insects. Stockport. 143-218. Moore, B. P. (1957). T h e British Carabidae (Coleoptera) part 2. T h e County Distribution of the Species. Ent. Gazette 8, 171-180. Morley, C. (1899). The Coleoptera of Suffolk. Plymouth. xiv and 113 pp., map. Morley, C. (1911). The Victoria History of the County of Suffolk, 1, 122128. Morley, C. (1915). The Coleoptera of Suffolk. First Supplement. Plymouth. iv and 12 pp. Morris, M. G. (1963). Notes on Huntingdonshire Weevils (Col. Curculionoidea) with special reference to Monks Wood and Wood Walton Fen National Nature Reserves. Ent. Gazette 14, 129-139. Morris, M. G. (1965). Notes on Huntingdonshire Weevils (Col. Curculionoidea) in 1963 and 1964 with special reference to the National Nature Reserves. Ent. Gazette 16, 105-113. Omer-Cooper, J. (19261. The Victoria History of the County of Huntingdon 1, 95-117. Omer-Cooper, J., Perkins, M. G. L., and Tottenham, C. E. (1928). T h e Coleoptera of Wicken Fen, 1 Introductory, 2 Geodephaga. Nat. Hist. Wicken Fen part 4, 267-297. Omer-Cooper, J. and Tottenham, C. E. (1932). 3 Hydradephaga, 4 Palpicornia, 5 Staphylinidae-Ipidae. Nat. Hist. Wicken Fen part 6, 489-538. Omer-Cooper, J. and Tottenham, C. E. (1934). Coleoptera taken in the air at Wicken Fen. Ent. mon. Mag. 70, 231-234. Tottenham, C. E. (1949). T h e Generic names of the British Staphylinidae with a check list of the species. Roy. Ent. Soc. Lond. The Genertc Names of British Insects part 9, 345-466. Various County Records, mainly by Claude Morley. Trans. Suffolk Nat. Soc. 1929-67, vols. 1-13. Various County Records, mainly by E. A. Ellis. Trans. Norfolk and Normch Nat. Soc. 1870-1967, vols. 1-21.

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